References
Sea Surface
Temperature
(SST, Ifaty)
Oxygen isotopes of coral 1660-1994 334 Zinke et al., 2022
Southern Annual
Mode Index (SAM)
Mid-latitude to polar
domain proxy records
1000-2007 1007 Abram et al.,
2014
Pacific Decadal
Oscillation (PDO)
Tree rings 993-1996 1003 MacDonald &
Case 2005
HadISST1.1 SST
dataset (Index
referring to IOD)
Calculated anomalous
SST gradient between the
western equatorial Indian
Ocean and the south-
eastern equatorial Indian
Ocean. Based on coral
isotopes and Ca/Mg
ratios.
1981-2010 29 Saji & Yamagata
2003
266
267 The identification of monotonic trends within the record was conducted using a non-
268 parametric Mann-Kendall trend test (Nasri & Modares, 2009; Pohlert, 2018) combined with a
269 Least Squares Regression to evaluate the rate of change in rainfall per year in the regional
270 records. In addition, gridded datasets downloaded from GPCC monthly total precipitation
271 from around Betioky at 2.5° x 2.5° resolution were compared to the composite record. For
272 correlations, significance at a level of 0.05 were accepted.
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273 3. RESULTS
274 The calibrated radiocarbon dates for the four trees suggest that they grew over the last 700 years
275 (1302 – 2013 CE) (Table SI1). The most parsimonious age models, that reconcile the AMS dates
276 and stable carbon isotope records, are shown in Fig. 2. All the trees show linear growth over time
277 except for GTR, which demonstrated a hiatus from 1500 CE to 1700 CE. This is not uncommon
278 in baobabs (Patrut et al., 2017). The age model assigned most of the AMS ages within 1-sigma
279 error of 68%.
280
281 Fig. 2: Age-models for (A) DFL, (B) DFS, (C) GTR with the hiatus indicated in dashed line, and
282 (D) TSP, based on 52 radiocarbon dates. The horizontal lines are the 1-sigma calibration
283 intervals for the radiocarbon dates. The bold line represents the age model that best intercepts the
284 1-sigma calibration range for the radiocarbon dates
285 The corrected δ13C time series from the four trees range between -26.2‰ (1400 CE) and -24.5‰
286 (1650 CE) with a mean of -25.3‰, and a variation of about 1.7‰ (Fig. 3). The trend analysis on
287 the composite δ13C records using the Mann-Kendall test shows that there is a marginal
288 decreasing trend of the isotope data over time (S=-5.07, p<0.01) with a difference of about -
289 0.7‰ between 1300 CE and 2013 CE. A least square linear regression is significant (F= 80.6, p
290 = 0.001) but with an R2 of 0.10. The change point analysis revealed a number of major shifts in
291 the mean values of the composite isotope data becoming either more positive or negative in the
292 time series (significant at 95%). These occur at approximately 1400 CE, 1480 CE, 1500 CE,
293 1630 CE, 1660 CE, 1820 CE, and 980 CE (Fig. 4).
294
295 Fig. 3: Corrected δ13C time series of four baobabs from southwest Madagascar with inverted y-
296 axis indicating drier (less negative isotope value) and wetter conditions (more negative isotope
297 value).
298
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299 Fig. 4: The composite record from four trees (black) with wetter periods (blue) and drier periods
300 (yellow). Error bars represent standard errors.
301 The isotope biweight mean composite and the GPCC monthly total precipitation both shows
302 similar trends over time. A linear regression of precipitation data from the GPCC monthly total
303 precipitation from 1900–present revealed a decreasing but not significant trend over time (β = -
304 0.67, p = 0.17), with year explaining only ~1.7% of the variance (R² = 0.017). Conversely, the
305 isotope data showed a small but significant increasing trend (β = +0.00112, p = 0.015) reflecting
306 decreasing rainfall, though the model explains less than 1% of variance (R² = 0.0099). These
307 results suggest subtle but differing temporal behaviours in the two proxies potentially associated
308 with the associated ages. Both records show a slight decrease in rainfall as reflected by the
309 precipitation value and the more positive isotope value around 1950 and then from 1970-1990
310 while more wetter periods are recorded around 1960 and 2000 (Fig. 5).
311
312 Fig. 5: Comparison of the baobab δ13C records with existing model datasets: Black indicates the
313 baobab δ13C composite records from 1900-2015 with linear regression indicated in grey. Blue
314 and dark blue shows GPCC total precipitation at 2.5° x 2.5° resolution from around Betioky
315 between 1900-2013 along with the associated linear model.
316 The correlation analysis of potential rainfall drivers in Southwest Madagascar has been
317 calculated (Table 3). The results diverse correlation values and significance (Table 3). There is a
318 negative correlation between SST and the isotope data an overall significant positive correlation
319 between SAM and the isotope records with recorded weak correlation since the 16th century.
320 Related to the Pacific Decadal Oscillation (PDO) reconstruction, no significant correlation has
321 been recorded during the entire period but during the LIA negative PDO anomalies are
322 associated with decreases in rainfall particularly between 1600 – 1750 CE (r=-0.30, p<0.001),
323 whereas in the record before and after this period, they are associated with increases in rainfall
324 (r=0.18, p=0.004 and r=0.18, p=0.001 respectively) (Fig. 7C).
325
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326 Table 3: Correlation between baobab isotope data and various environmental indices.
327 Pearson correlation coefficients (r) and associated p-values are shown. Sample size (n) indicates
328 the number of data points compared for each dataset.
Dataset Compared with Baobab
Isotope Data
Time Period
(AD)
Sample Size
(n)
Pearson
Correlation
(r)
p-value
Sea Surface Temperature (SST,
Ifaty) 1660–1994 334 –0.22* < 0.001*
Southern Annual Mode Index
(SAM) 1000–2007 1007 0.16* 0.05
HadISST1.1 SST dataset (IOD
Index) 1981–2010 29 –0.06 > 0.05
329 Note: * indicates significance at α = 0.05.*
330 4. DISCUSSION
331 4.1. Rainfall record of southwest Madagascar for the last 700 years
332 The similarities between the δ13C records and the GPCC precipitation data meets the theoretical
333 expectation, and reaffirms the results obtained for baobab isotope controls on the African
334 mainland (Woodborne et al., 2015; 2016; Fig. 5). The baobab δ13C record can thus be interpreted
335 as a proxy for local effective rainfall in southwest Madagascar, reflecting decadal to centennial
336 variability in the last 700 years. Accordingly, lower δ13C values indicate wetter periods while
337 more positive isotope ratios correlate to drier conditions.
338 The chronology of the four baobabs ranges from 1300 CE until 2013 with a hiatus in the GTR
339 core from 1500 CE to 1700 CE. Punctuated growth models have been noted in other baobabs
340 (Patrut et al., 2017) and may arise because of attenuated growth rates over time, or possibly due
341 to lobate development of the stem resulting in the reallocation of resources to another part of the
342 tree. The GTR baobab was collected near the River Mangoky in southwest Madagascar where a
343 sediment core was taken from nearby Lake Tsizavatsy. The age-depth model of the Tsizavatsy
344 core shows a hiatus from 1400 to 1900, suggested to be associated with a regional drought event
345 (Razanatsoa et al., 2021). Although the other trees show positive isotope excursions (dry events)
346 during this period, there are oscillations of wetter conditions covered by the present records (Fig.
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347 3). Notwithstanding the gap, the record from GTR prior to and after the hiatus was combined
348 with the composite record of all trees to provide a full rainfall proxy record for the last 700 years
349 for southwest Madagascar covering the Little Ice Age (LIA; Lechleitner et al., 2017; Putnam &
350 Broecker, 2017) and the Anthropocene.
351 Synchronicity in the baobab δ13C records from southwest Madagascar demonstrate regional
352 variability with a succession of wet and dry cycles on the scale of decades to centuries (Fig. 4).
353 The rainfall in the region was suggested to be variable with drought events recorded in
354 Southwest Madagascar prior 1000 CE based on stalagmite records (Faina et al., 2021) with
355 drying trends being recorded on the isotope tree records since 1300 CE. Both records, baobab
356 δ13C and stalagmite δ¹⁸O show similar trends and patterns demonstrating high agreement in the
357 variability of rainfall over time despite a weak negative Pearson correlation between the two
358 variables (See Table 3). This non correlation could be explained by the uncertainties associated
359 with the two age models. Before 1700 CE, both records show different trends with discrepancies
360 around mid-17th century where the tree record showed drier conditions while the stalagmites is
361 showing wetter conditions. Post 1700 CE similarities between the stalagmite 𝛿18O and 𝛿13C tree
362 records are recorded with wet condition around 1700-1800 CE, dry condition 1800-1900 CE and
363 a wetter condition again between 1900-2000 CE. Very recently (post-1950 CE), both records
364 reflected a severe drought (Fig. 6).
365 Fig. 6: Comparison between baobab δ13C records from tree rings in southwest Madagascar
366 (black) with stalagmite δ18O records from Asafora cave from the southwest coast and just
367 southeast of the Velondriake Marine Reserve (green, from Faina et al., 2021).
368 The composite baobab record shows increasing δ13C values indicating marginal drying at the
369 locations of the trees over the past 700 years, corresponding with previous findings suggesting
370 increasing aridity since 1000 CE (e.g. Burney, 1993; Burns et al., 2016; Virah-Sawmy et al.,
371 2016, Razanatsoa et al., 2021). Pollen records of the last millennia show a synchronous
372 desiccation from various regions in Madagascar including the southwest (Burney, 1993; Burns et
373 al., 2016; Virah-Sawmy et al., 2016) which agrees with reduced length of wet periods in the tree
374 records compared to regional records.
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375 4.2. Synoptic drivers of rainfall in southwest Madagascar
376 When compared with other baobab δ13C records from South Africa (Woodborne et al., 2015;
377 2016), the rainfall patterns for southwest Madagascar and the South African summer rainfall
378 zone are in phase for most of the last 700 years. Wet (dry) periods in the South African Pafuri
379 and Mapungubwe records (22 °S) corresponded to similar wet (dry) periods from southwest
380 Madagascar. This suggests that southwest Madagascar rainfall responds to similar drivers of the
381 summer rainfall zone in southern Africa, including Agulhas Current sea-surface temperature
382 variations regulated by the East-West displacement of the TTTs (Woodborne et al., 2016) in
383 addition to the regulatory effect of the island’s mountains (Donque, 1975; Jury & Huang, 2004).
384 However, further investigation of the various drivers of rainfall is required to provide more
385 understanding of local and regional temporal changes.
386 4.2.1. ITCZ and SAM modulated rainfall during Early Little Ice Age between 1370
387 and 1500
388 The composite baobab record from southwest Madagascar shows a wet period between
389 approximately 1370 – 1500 CE (Fig. 4) which coincides with the second phase 1495 – 1833 CE
390 of a wet-neutral-wet cycle recorded in northwest Madagascar (Scroxton et al., 2017). The
391 movement of the ITCZ has a significant impact on rainfall variability in Madagascar (Haug et
392 al., 2001; Liu et al., 2003; Verschuren et al., 2000; Schneider et al., 2014). A southward shift was
393 recorded at the beginning of the Little Ice Age around 1300 CE (Chiang & Bitz, 2005; Broccoli
394 et al., 2006; Lechleitner et al., 2017; Putnam & Broecker, 2017) and this may have resulted in
395 increased rainfall for southwestern Madagascar. Wetter conditions are also evident in several
396 East African sediment records during this period, including Lake Chilwa (Crossley et al., 1984),
397 Lake Malawi (Johnson et al., 2001), Lake Massoko (Barker et al., 2000), Lake Tanganyika (Alin
398 & Cohen 2003), Lake Victoria (Stager et al., 2005), Lake Naivasha (Verschuren et al., 2000;
399 Tyson et al., 2001; Tierney et al., 2013) suggesting a common driver of rainfall, most likely the
400 southwards movement of the ITCZ.
401 The wetter conditions from 1370 – 1500 CE in southwestern Madagascar occur when the
402 Southern Annular Mode (SAM) was at its most extreme negative phase in the fifteenth century
403 (Abram et al., 2014; Fig. 7A). Negative SAM indices imply an expansion of the westerly
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404 circumpolar vortex, which has significant impacts on temperature and precipitation over all four
405 Southern Hemisphere continents (Gillett et al., 2006), and particularly over Africa south of 25°S,
406 where an increase in precipitation is associated with the northward migration of the westerlies
407 during the austral winter. The response is most pronounced along the east coast, where it is
408 associated with anomalous easterly winds advecting more moisture off the SWIO (Gillett et al.,
409 2006). No significant response to SAM has been reported for Madagascar on the basis of
410 instrumental data (Gillett et al., 2006) although heavy rainfall events have been associated to
411 atmospheric circulation displays a Southern Annular Mode-like pattern throughout the
412 hemisphere (Randriamahefasoa & Reason, 2017). The relationship between SAM and rainfall in
413 the baobab record presented here is not consistent with an overall significant positive correlation
414 (r=0.16, p<0.001). However, since the 16th century the relationship is weak and non-significant
415 suggesting that the southward contraction of the westerly winds during the positive SAM reduces
416 their influence on the region. When the westerlies migrate southward during a positive SAM
417 phase, they cease to be a driver of rainfall, and decadal to centennial rainfall variability responds
418 to other forcing. The lack of consistent correlation throughout the records also supports recent
419 findings related to the lack of a forced response in SAM variability prior to the 20th century
420 (King et al 2023).
421 Our composite record suggests that rainfall responds to subtropical forcing during extreme
422 negative SAM phases as the westerly winds migrate northwards bringing wetter condition to the
423 subtropics including southern Madagascar (Fig. 7A), but otherwise it responds to tropical forcing
424 determined by the position of the ITCZ during the austral summers. Evidence of this mechanism
425 operating during glacial periods is derived from marine records and model analysis of the late
426 Glacial maximum over southern Africa (Anderson et al., 2009; Sigman et al., 2010; Miller et al.,
427 2019b; Engelbrecht et al., 2019; Hahn et al., 2021). The mechanism may explain the rainfall
428 maximum during the 14th and 15th centuries, but further research is needed to elucidate the
429 influence of the SAM on southern African climate during the Little Ice Age.
430 Fig. 7: Comparison of the baobab δ13C records (black) with (A) the Southern Annual Mode
431 (SAM) index (orange) (Abram et al. 2014), (B) the Pacific Decadal Oscillation (PDO) index
432 (blue) (MacDonald & Case 2005), (C) Sea Surface Temperature (SST) from Ifaty, southwest
433 Madagascar (grey) (Zinke et al. 2022), and (D) HadISST1.1 SST dataset that refers to Indian
434 Ocean Dipole variability (red, Saji & Yamagata 2003)
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435 4.2.2. Regional and localised rainfall drivers during the LIA 1600-1750
436 Wet conditions during the 14th and 15th centuries are followed by extremely dry conditions 1600
437 – 1750 CE in southwest Madagascar. This period is characterised by a positive phase of the
438 SAM that commences at 1500 CE (Abram et al., 2014) implying reduced effect of the westerlies.
439 It is also a period of reduced sunspot activity known as the Maunder Minimum (1645 – 1715)
440 (Mann et al., 2009; Scroxton et al., 2017) with decreased global temperatures at the maximum of
441 the Little Ice Age (LIA; Tyson et al., 2001). It has been speculated that there was a southward
442 migration of the ITCZ during the maximum of the LIA (Jury & Huang, 2004; Russel et al., 2007;
443 Tadross et al., 2008; Voarintsoa et al., 2017). Our results show drier condition during this period
444 starting around 1600 CE and has also been experienced in northwest of Madagascar around 1700
445 CE (Scroxton et al., 2017). Dry periods were also experienced across the African continents
446 including East Africa (Russell & Johnson, 2007; Tierney et al., 2013), and the southern African
447 summer rainfall area (Huffman, 2004; PAGES 2k Consortium, 2013; Macron et al., 2014;
448 Chevalier & Chase, 2015; Huffman & Woodborne, 2016; Woodborne et al., 2016). Pollen
449 evidence from the Lake Longiza in southwest Madagascar suggested an increase in grass and
450 decrease in trees and shrubs such as Arecaceae, Pandanus, and Acacia potentially associated
451 with the drying in the region (Matsumoto & Burney, 1994; Razanatsoa et al., 2022). Lake
452 Tsizavatsy, from the same region, showed a hiatus in its sediment deposition between 1400 –
453 1900 CE (Razanatsoa et al., 2021) suggesting the drying of the lake during this period, which is
454 consistent with the tree records. The evidence of dry conditions from records across southern
455 Africa does not support southward migration of the ITCZ during this period and if the migration
456 occurred, there might have been other drivers that inhibited its effect leading to a decrease in
457 rainfall during this period.
458 The Pacific Decadal Oscillation (PDO) is linked to ENSO in relation to drought patterns across
459 Africa through teleconnections that influence the longitudinal position of climate systems. The
460 (PDO) reconstruction shows that during the LIA negative PDO anomalies are associated with
461 decreases in rainfall particularly between 1600 – 1750 CE (r=-0.30, p<0.001), whereas in the
462 record before and after this period, they are associated with increases in rainfall (r=0.18, p=0.004
463 and r=0.18, p=0.001 respectively) (Fig. 7C). Despite the change in the sign of the correlation,
464 this evidence suggests that climate forcing is dominated by tropical forcing. (Thompson et al.,
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465 2003; MacDonald & Case 2005; Hoell et al., 2017). Changes in the TTTs position which tend to
466 propagate eastward, from southern Africa to the Mozambique Channel and southern Madagascar
467 is known to have a strong influence on intra-seasonal and even interannual rainfall variability in
468 the region (Macron et al., 2014). This has particularly been suggested to be dictated by the
469 migration of the ITCZ (Chase & Meadows, 2007; Sachs et al., 2009) and to increase during La
470 Nina conditions (Manhique et al., 2011; Ratna et al., 2012; Macron et al., 2014). Moreover, its
471 persistence was suggested to be maintained by variation of SST anomalies over the Agulhas
472 Current (Manhique et al., 2011; Vigaud et al., 2007). Comparison of the baobab δ13C records
473 from 1660 – 1994 CE with the 300-year Agulhas Current Sea surface temperature (SST) record
474 from Ifaty in southwest Madagascar (Zinke et al., 2014) shows a significant negative correlation
475 (r=-0.22, p<0.001). Positive (negative) SST corresponds with higher (lower) rainfall in the
476 baobab record (Fig. 7B). The coolest oceanic temperatures in the coral record, with anomalies of
477 -0.3 – -0.5 °C between 1675 – 1760 CE, correspond to the driest period in southwest
478 Madagascar. Similar patterns of rainfall were noted in the summer rainfall area of the adjacent
479 African mainland (Woodborne et al., 2015). Variation in SST in the western Indian Ocean are
480 determinant in the IOD with a suggested negative relationship with eastern African rainfall
481 responses (Hoell et al., 2017; Taylor et al., 2021).
482 4.2.3. Mixed effect of changes in ITCZ position, human land-use and climate change
483 from 1750 – 2013 CE
484 Around 1750 CE until early 1800 CE, at the end of the LIA, there is a relatively wet period
485 recorded in the baobab δ13C data. A relatively dry period after 1860 CE is similar to conditions
486 experienced over the summer rainfall zone in southern Africa. These periods coincided with
487 more extreme ENSO warm phases (Nash, 2017) with the warmest period in the Agulhas SST
488 record between 1880 CE and 1900 CE and a northward migration of the ITCZ (Zinke et al.,
489 2014; Railsback et al., 2018).
490 The comparison of the composite baobab δ13C record with the Dipole Mode Index (DMI) that
491 were used as an index of the IOD (1870 – 2013) shows very low to no correlation (Table 3, Fig.
492 7D) but with a noticeable positive but not significant correlation since 1980 CE (r= 0.27, p=
493 0.08). The effect that the IOD has on equatorial climate forcing is similar to the ENSO or PDO
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494 effects as it is driven by an equatorial SST differential across the Indian Ocean while ENSO is
495 driven by a gradient in the Pacific Ocean. The effect in the subtropical region of southwestern
496 Madagascar appears to be dominated by the Pacific Ocean influences on global climate
497 (ENSO/PDO) which is influential in the latitudinal position of the TTTs system.
498 Around 1950 CE, conditions in southwest Madagascar are as wet as at any time in the record.
499 This corresponds to a positive SST anomaly, positive phase of IOD and a negative PDO phase.
500 Despite suggested changes in the impact of ENSO cycles on the SST in the region of the SWIO
501 since 1970 (Zinke et al., 2014), our results show typical PDO/rainfall phasing with more
502 negative PDO corresponding to high rainfall while the inverse is not always true. Records
503 suggest that the IOD intensified following the onset of global warming during the 20th century
504 along with forced response of SAM (Abram et al., 2008; Namakura et al., 2009; Watanabe et al.,
505 2019; King et al 2023), with increased evidence of human induced climate change (IPCC, 2021),
506 evidence of increased river runoff and shifts in human land-use through slash and burn
507 agriculture were recorded in coral records from eastern Madagascar (Grove et al., 2013). Pollen
508 records from the region suggest a decrease in the tree component including Arecaceae coinciding
509 with an increase in Poaceae and pioneer taxa such as Asteraceae mostly likely associated with
510 tree cutting associated with agriculture expansion (Razanatsoa et al., 2022). These suggest that
511 changes in rainfall in the region led to changes in land-use. There is a return to drier conditions
512 around 1980 called “belt of iron” also recorded in historical records peaking at the beginning of
513 the 1990s (Von Heland & Folke, 2014). This was followed by a trend towards wet conditions in
514 the past 20 years similar the instrumental record (Tadross et al., 2008).
515 4.3. Implications for future climate change risk and adaptation
516 The last 300 years of the baobab composite record show the interacting global, regional and local
517 drivers that have influenced rainfall variability in southwest Madagascar. The dominant effect of
518 the position of the ITCZ during the austral summer is evident, and this is troubling in the context
519 of forecast climate change. The position and zone of the ITCZ is predicted to narrow with a
520 northward shift over eastern Africa and the Indian Ocean and a southward shift in the eastern
521 Pacific and Atlantic oceans by 2100, which would severely reduce rainfall in the region
522 (Mamalakis et al., 2021). In terms of climate risk and adaptation, southwest Madagascar will
523 likely experience a drier climate with more frequent and prolonged droughts as already predicted
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524 in the IPCC 6th assessment report (IPCC, 2021). Climatic factors are an important driver of
525 economic, environmental and societal decisions (Boutin & Smit, 2016) and would be crucial for
526 the future of these dry areas. Indeed, the population are dependent on rainfall as a source of
527 water and for agriculture due to the lack of infrastructure and the limited permanent water ponds
528 (Hänke et al., 2017; Carriere et al., 2018). The effect of drought and lack of rainfall has already
529 led the Mikea forager communities to diversify their livelihoods with seasonal agriculture to
530 ensure food security (Razanatsoa et al., 2021). Some adaptations that have been established
531 elsewhere and could be conducted in the region include the introduction of new drought resistant
532 crops, (e.g. Thomas et al., 2007; Yaro et al., 2014). Multiple species livestock herding with cattle
533 and goat pastoralism (Kaufmann & Tsirahamba, 2006; Hänke & Barkmann, 2017) and livelihood
534 diversification including work that is not farming (e.g. crafts and services for local markets) were
535 suggested to be a major adaptive strategy under drying conditions in a short and long term and
536 buffer livelihoods in the face of environmental change (e.g. Kuiper et al., 2007; Tambo &
537 Abdoulaye, 2013). Further understanding of the effect of future climate on these populations and
538 their surrounding environment is critical in planning strategies of adaptation in terms of
539 livelihoods but also water provision also in the coming years.
540 5. CONCLUSION
541 Baobab δ13C data from southwest Madagascar are a proxy for changing rainfall over the last 700
542 years. The inferred wettest period was between 1370 – 1500 CE while the driest period occurred
543 between 1600 – 1750 CE. High centennial variability was recorded with a decreasing rainfall
544 trend and reducing duration of wet periods over time. The comparison of the records with
545 existing records of rainfall drivers at local, regional, and global scales shows that the baobab
546 rainfall proxy record is not dominated by the influence of any single forcing over the entire
547 record. The westerlies may play a role during extreme negative phases of the SAM, while
548 latitudinal shifts of the ITCZ are the dominant low frequency driver of rainfall. At a more local
549 level, the role of SST seems to dominate variability possibly through longitudinal influences on
550 the position of the TTTs system which is also influenced by the PDO/ENSO system. Localised
551 climate forcing in relation to the Southwest Madagascar Coastal Current (SMACC) within the
552 greater Agulhas Current system has been suggested (Ramanantsoa et al., 2018). What emerges
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553 from comparisons with other rainfall proxy records on the island of Madagascar, and from the
554 adjacent African mainland, is that the temporal trends are not consistent, probably reflecting the
555 contrasting dominance of different drivers in different regions. A rainfall dipole exists between
556 southern Africa and Madagascar (Jury et al., 2015; 2016; Woodborne et al., 2016; Barimalala et
557 al., 2018) with increases in precipitation over southern Africa extending from Mozambique to
558 Angola coincident with a decrease in rainfall most of Madagascar (Barimalala et al., 2018) but
559 not in the southwest region. The mountains that extend from the north to the south of
560 Madagascar (>1500 m elevation) reduce the direct transport of moisture from the Indian Ocean
561 toward southern Africa (Barimalala et al., 2018) and southwest Madagascar. The Agulhas
562 current SST forcing of Madagascar rainfall is opposite to that in southern African where negative
563 SST anomalies were associated with wetter conditions over the southern African interior
564 (Woodborne et al., 2015). These contradictions suggest that the variation in rainfall is not a
565 simple intensification or weakening of the existing climate patterns, but rather a response in the
566 synoptic systems to tropical (ENSO/PDO), extratropical (SAM), and localised (SST) forcing.
567 Why some forcing appears to dominate at certain periods and not at others is unclear, and the
568 evidence presented here suggests that synergistic effects might be explored in global climate
569 models.
570 The potential effect of climate change and land-use change were also recorded at the near present
571 period, as well as possible effects of SAM if there is a northward migration of westerlies similar
572 to what happened around 1300 CE. The data generated here provide the opportunity to unravel
573 the relative importance and interaction between global, regional, and local drivers across the
574 southern and eastern African region. These findings are crucial in the simulations of rainfall
575 projections to help evaluate the impacts and trends of migration of the westerlies and
576 anthropogenic induced climate change on the African continents that are not fully understood.
577 For southwest Madagascar with an expected drier climate and increasing occurrence of severe
578 drought conditions predicted for the near future, understanding the risks, and establishing
579 adaptation strategies particularly in terms of livelihood could avoid disastrous famine.
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1. ACKNOWLEDGEMENT
We would like to thank the Ministry of Environment and Sustainable Development of
Madagascar and Madagascar National Park for providing the permission for the field campaign,
sampling and exportation. We acknowledge ESSA-Forêts Mention Foresterie et Environnement
de l’Ecole Supérieure des Sciences Agronomiques, Université d’Antananarivo –
MADAGASCAR for collaborating with the obtention of the research permit. We also
acknowledge all the field assistants that have participated in retrieving the cores and Tsilavo
Razafimanantsoa for his input on the map.
1. DATA REFERENCES
Zinke, J., B. Loveday, C. Reason, W.-C. Dullo, and D. Kroon. (2014). Madagascar corals track
sea surface temperature variability in the Agulhas Current core region over the past 334
years. Scientific Reports, 4, 4393. doi: 10.1038/srep04393. NOAA's National Centers for
Environmental Information (NCEI)
Abram, N. J., Mulvaney, R., Vimeux, F., Phipps, S. J., Turner, J., & England, M. H. (2014).
Evolution of the Southern Annular Mode during the past millennium. Nature Climate
Change, 4(7), 564–569. doi.org/10.1038/nclimate2235. NOAA's National Centers for
Environmental Information (NCEI)
Jinbao Li, Shang-Ping Xie, Edward R. Cook, Gang Huang, Rosanne D'Arrigo, Fei Liu, Jian Ma,
and Xiao-Tong Zheng. 2011. Interdecadal modulation of El Niño amplitude during the past
millennium. Nature Climate Change. 1(2) 114-118. doi: 10.1038/nclimate1086. NOAA's
National Centers for Environmental Information (NCEI)
Scroxton, N., Burns, S. J., Mcgee, D., Hardt, B., Godfrey, L. R., Ranivoharimanana, L., & Faina,
P. (2017). Hemispherically in-phase precipitation variability over the last 1700 years in a
Madagascar speleothem record. 164. doi.org/10.1016/j.quascirev.2017.03.017. National
Centers for Environmental Information (NCEI)
.CC-BY 4.0 International licensemade available under a
(which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is
The copyright holder for this preprintthis version posted August 20, 2025. ; https://doi.org/10.1101/2025.08.15.670475doi: bioRxiv preprint
MacDonald & Case. 2005. Pacific Decadal Oscillation Reconstruction for the Past Millennium.
GRL 32, L08703. doi:10.1029/2005GL022478. National Centers for Environmental
Information (NCEI)
Saji, N.H., & Yamagata, T., (2003). Possible impacts of Indian Ocean Dipole mode events on
global climate. CLIMATE RES, 25 (2): 151-169.
https://psl.noaa.gov/gcos_wgsp/Timeseries/DMI/
2. DATA AVAILABILITY
Datasets related to this article can be found at 10.25375/uct.16590035 on Zivahub, an open-
source online data repository hosted by the University of Cape Town. The DOI will become
active upon publication of the manuscript.
3. REFERENCES
Abram, N. J., Gagan, M. K., Cole, J. E., Hantoro, W. S. & Mudelsee, M. (2008). Recent
intensification of tropical climate variability in the Indian Ocean. Nature Geoscience 1,
849–853.
https://doi.org/10.1038/ngeo357
Abram, N. J., Mulvaney, R., Vimeux, F., Phipps, S. J., Turner, J., & England, M. H. (2014).
Evolution of the Southern Annular Mode during the past millennium. Nature Climate
Change, 4(7), 564–569. https://doi.org/10.1038/nclimate2235
Alin, S. R., & Cohen, A. S. (2003). Lake-level history of Lake Tanganyika, East Africa, for the
past 2500 years based on ostracode-inferred water-depth reconstruction. Palaeogeography,
Palaeoclimatology, Palaeoecology, 199(1–2), 31–49. https://doi.org/10.1016/S0031-
0182(03)00484-X
Anderson, R. F., Ali, S., Bradtmiller, L. I., Nielsen, S. H. H., Fleisher, M. Q., Anderson, B. E.,
and Burckle, L. H. (2009). Wind-Driven Upwelling in the Southern Ocean and the
Deglacial Rise in Atmospheric CO2, Science. 323, 1443–1448,
https://doi.org/10.1126/science.1167441.
.CC-BY 4.0 International licensemade available under a
(which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is
The copyright holder for this preprintthis version posted August 20, 2025. ; https://doi.org/10.1101/2025.08.15.670475doi: bioRxiv preprint
Baptista, D. M. S., Farid, M., Fayad, D., Kemoe, L., Lanci, L. S., Mitra, P., Muehlschlegel, T. S.,
Okou, C., Spray, J. A., Tuitoek, K., & Unsal, F. D. (2022). Climate change and chronic
food insecurity in sub-Saharan Africa. International Monetary Fund Library, 2022(16), 1.
https://doi.org/10.5089/9798400218507.087
Barimalala, R., Desbiolles, F., Blamey, R. C., & Reason, C. (2018). Madagascar Influence on the
South Indian Ocean Convergence Zone, the Mozambique Channel Trough and Southern
African Rainfall. Geophysical Research Letters, 45(20), 11,380-11,389.
https://doi.org/10.1029/2018GL079964
Barimalala, R., Raholijao, N., Pokam, W., & Reason, C. J. C. (2021). Potential impacts of 1.5
°C, 2 °C global warming levels on temperature and rainfall over Madagascar.
Environmental Research Letters, 16(4). https://doi.org/10.1088/1748-9326/abeb34
Barker, P., Telford, R., Merdaci, O., Williamson, D., Taieb, M., Vincens, A., & Gibert, E.
(2000). The sensitivity of a Tanzanian crater lake to catastrophic tephra input and four
millennia of climate change. Holocene, 10(3), 303–310.
https://doi.org/10.1191/095968300672848582
Baudoin, M.A., Vogel, C., Nortje, K., Naik, M. (2017). Living with drought in South Africa:
lessons learnt from the recent El Niño drought period. International Journal of Disaster
Risk Reduction., 23 , 128-137. https://doi.org/10.1016/j.ijdrr.2017.05.005
Baum, D. A., Bombacaceae, A., & Baum, D. A. (1995). A Systematic Revision of Adansonia
(Bombacaceae) 82(3), 440–471. https://doi.org/10.2307/2399893
Baum, D. A., Small, R. L., & Wendel, J. F. (1998). Biogeography and flora evolution of Baobabs
(Adansonia, Bombacaceae) as infered from multiple data sets. Systematic Biology, 47(2),
181–207.
https://doi.org/10.1080/106351598260879
Beck, H. E., Wood, E. F., Pan, M., Fisher, C. K., Miralles, D. M., van Dijk, A. I. J. M., McVicar,
T. R., and Adler, R. F. MSWEP V2 global 3-hourly 0.1° precipitation: methodology and
quantitative assessment Bulletin of the American Meteorological Society 100(3), 473–500,
2019
.CC-BY 4.0 International licensemade available under a
(which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is
The copyright holder for this preprintthis version posted August 20, 2025. ; https://doi.org/10.1101/2025.08.15.670475doi: bioRxiv preprint
Belmecheri, S., & Lavergne, A. (2020). Compiled records of atmospheric CO2 concentrations
and stable carbon isotopes to reconstruct climate and derive plant ecophysiological indices
from tree rings. Dendrochronologia, 63(August), 125748.
https://doi.org/10.1016/j.dendro.2020.125748
Broccoli, A. J., Dahl, K. A., & Stouffer, R. J. (2006). Response of the ITCZ to Northern
Hemisphere cooling. Geophysical Research Letters, 33(1), 1–4.
https://doi.org/10.1029/2005GL024546
Burney, D. (1993). Late Holocene Environmental changes in arid southwestern Madagascar.
Quaternary Research, 40, 98–106. https://doi.org/10.1006/qres.1993.1060
Burns, S. J., Godfrey, L. R., Faina, P., McGee, D., Hardt, B., Ranivoharimanana, L., &
Randrianasy, J. (2016). Rapid human-induced landscape transformation in Madagascar at
the end of the first millennium of the Common Era. Quaternary Science Reviews, 134, 92–
99. https://doi.org/10.1016/j.quascirev.2016.01.007
Carrière, S.D., Chalikakis, K., Ollivier, C. et al. (2018). Sustainable groundwater resources
exploration and management in a complex geological setting as part of a humanitarian
project (Mahafaly Plateau, Madagascar). Environmental Earth Science 77, 734
https://doi.org/10.1007/s12665-018-7909-1
Chase, B. M., & Meadows, M. E. (2007). Late Quaternary dynamics of southern Africa’s winter
rainfall zone. Earth-Science Reviews, 84(3–4), 103–138.
https://doi.org/10.1016/j.earscirev.2007.06.002
Chevalier, M., & Chase, B. M. (2015). Southeast African records reveal a coherent shift from
high- to low- latitude forcing mechanisms along the east African margin across last glacial
e interglacial transition. Quaternary Science Reviews, 125, 117–130.
https://doi.org/10.1016/j.quascirev.2015.07.009
.CC-BY 4.0 International licensemade available under a
(which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is
The copyright holder for this preprintthis version posted August 20, 2025. ; https://doi.org/10.1101/2025.08.15.670475doi: bioRxiv preprint
Chiang, J. C. H., & Bitz, C. M. (2005). Influence of high latitude ice cover on the marine
Intertropical Convergence Zone. Climate Dynamics, 25(5), 477–496.
https://doi.org/10.1007/s00382-005-0040-5
Christensen J.H., Hewitson B., Busuioc A., Chen A., Gao X., Held I., Jones R., Kolli R.K.,
Kwon W-T., Laprise R., Magana Rueda V., Mearns L., Menendez C.G., Raisanen J., Rinke
A., Sarr A., Whetton P. (2007). Regional climate projections. In: Solomon S., Qin D.,
Manning M., Chen Z., Marquis M., Averyt A.B., Tignor M., Miller H.L. (eds) Climate
change 2007: the physical science basis. Contribution of Working Group I to the Fourth
Assessment Report of the Inter-governmental Panel on Climate Change. Cambridge
University Press, Cambridge
Connolly-Boutin, L. & Smit, B. (2016). Climate change, food security, and livelihoods in sub-
Saharan Africa. Regional Environmental Change, 16: 385-399.
https://doi.org/10.1007/s10113-015-0761-x
Cook, E. R., Palmer, J. G., Cook, B. I., Hogg, A., & D’Arrigo, R. D. (2002). A multi-millennial
palaeoclimatic resource from Lagarostrobos colensoi tree-rings at Oroko Swamp, New
Zealand. Global and Planetary Change, 33(3–4), 209–220. https://doi.org/10.1016/S0921-
8181(02)00078-4
Crossley, R., Davison-Hirschmann, S., Owen, R.B., Shaw ,P.A. (1984). Lake level fluctuations
during the last 2000 years in Malawi. J. Vogel (Ed.), Late Cainozoic Palaeoclimates of the
Southern Hemisphere, A.A. Balkema, Rotterdam pp. 305-316. http://pascal-
francis.inist.fr/vibad/index.php?action=getRecordDetail&idt=7268013
Donque, G. (1972). The climatology of Madagascar. In: Biogeography and ecology of
Madagascar. R. Battistini and G. Richard-Vindard (eds). Junk, The Hague. 87–144.
https://doi.org/10.1007/978-94-015-7159-3_3
Engelbrecht, F. A., Marean, C. W., Cowling, R. M., Engelbrecht, C. J., Neumann, F. H., Scott,
L., Nkoana, R., O’Neal, D., Fisher, E., Shook, E., Franklin, J., Thatcher, M., McGregor, J.
L., Van der Merwe, J., Dedekind, Z., & Difford, M. (2019). Downscaling Last Glacial
.CC-BY 4.0 International licensemade available under a
(which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is
The copyright holder for this preprintthis version posted August 20, 2025. ; https://doi.org/10.1101/2025.08.15.670475doi: bioRxiv preprint
Maximum climate over southern Africa. Quaternary Science Reviews, 226, 105879.
https://doi.org/10.1016/j.quascirev.2019.105879
Faina, P., Burns, S. J., Godfrey, L.R., Crowley, B.E., Scroxton, N., McGee, D., Sutherland,
M.R., & Ranivoharimanana, L. Comparing the paleoclimates of northwestern and
southwestern Madagascar during the late Holocene: Implications for the role of climate in
megafaunal extinction. Malagasy nature, 15 (). Retrieved from
https://par.nsf.gov/biblio/10316730.
Farquhar, G., O’Leary, M. & Berry, J. (1982). On the Relationship between Carbon Isotope
Discrimination and the Intercellular Carbon Dioxide Concentration in Leaves. Australian
Journal of Plant Physiology. 9(2), 121-137. https://doi.org/10.1071/PP9820121
Ganzhorn, J. U. (1995). Cyclones over Madagascar: fate or fortune? Ambio. 24,124–125.
http://www.jstor.org/stable/4314308
Gillett, N. P., Kell, T. D., & Jones, P. D. (2006). Regional climate impacts of the Southern
Annular Mode. Geophysical Research Letters, 33(23), 1–4.
https://doi.org/10.1029/2006GL027721
Grove, C. A., Zinke, J., Peeters, F., Park, W., Scheufen, T., Kasper, S., Randriamanantsoa, B.,
McCulloch, M. T., & Brummer, G. J. A. (2013). Madagascar corals reveal a multidecadal
signature of rainfall and river runoff since 1708. Climate of the Past, 9(2), 641–656.
https://doi.org/10.5194/cp-9-641-2013
Hahn, A., Schefuß, E., Groeneveld, J., Miller, C., & Zabel, M. (2021). Glacial to interglacial
climate variability in the southeastern African subtropics (25–20° S). Climate of the Past,
17, 345–360. https://doi.org/10.5194/cp-2019-158
Hajdas, I., Hendriks, L., Fontana, A., & Monegato, G. (2017). Evaluation of Preparation
Methods
in Radiocarbon Dating of Old Wood. Radiocarbon, 59(3), 727–737.
https://doi.org/10.1017/RDC.2016.98
.CC-BY 4.0 International licensemade available under a
(which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is
The copyright holder for this preprintthis version posted August 20, 2025. ; https://doi.org/10.1101/2025.08.15.670475doi: bioRxiv preprint
Hall, G., Woodborne, S., & Pienaar, M. (2009). Rainfall control of the δ 13 C ratios of
Mimusops caffra from KwaZulu-Natal, South Africa. The Holocene, 19(2), 251–260.
https://doi.org/10.1177/0959683608100569
Hall, G., Woodborne, S., & Scholes, M. (2008). Stable carbon isotope ratios from archaeological
charcoal as palaeoenvironmental indicators. 247, 384–400.
https://doi.org/10.1016/j.chemgeo.2007.11.001
Hänke, H. & Barkmann, J. (2017). Insurance Function of Livestock: Farmer’s Coping Capacity
with Regional Droughts in South-Western Madagascar. World Development. 96: 264–275.
https://doi.org/10.1016/j.worlddev.2017.03.011
Hänke, H., Barkmann, J., Coral, C., Enforskaustky, E., & Marggraf, R. (2017). Social-ecological
traps hinder rural development in Southwestern Madagascar. Ecology and Society, 22(1).
https://doi.org/10.5751/ES-09130-220142
Haug, G., Hughen, K., Sigman, D., Peterson, L., & Ro, U. (2001). Southward Migration of the
ITCZ holocene. Science, 293(August), 1304–1309. https://doi.org/10.1126/science.1059725
Helama, S., Timonen, M., Lindholm, M., Meriläinen, J., & Eronen, M. (2005). Extracting long-
period climate fluctuations from tree-ring chronologies over timescales of centuries to
millennia. International Journal of Climatology, 25(13), 1767–1779.
https://doi.org/10.1002/joc.1215
Ho, C. H., Kim, J. H., Jeong, J. H., Kim, H. S., & Chen, D. L. (2006). Variation of tropical
cyclone activity in the South Indian Ocean: El Nino-Southern Oscillation and Madden-
Julian Oscillation effects. Journal of Geophysical Research-Atmospheres, 111(D22),
D22101. https://doi.org/Artn D22101\nDoi 10.1029/2006jd007289
Hoell, A., Funk, C., Zinke, J., & Harrison, L. (2017). Modulation of the Southern Africa
precipitation response to the El Niño Southern Oscillation by the subtropical Indian Ocean
Dipole. Climate Dynamics, 48(7–8), 2529–2540. https://doi.org/10.1007/s00382-016-
3220-6
.CC-BY 4.0 International licensemade available under a
(which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is
The copyright holder for this preprintthis version posted August 20, 2025. ; https://doi.org/10.1101/2025.08.15.670475doi: bioRxiv preprint
Hogg A.G., Hua Q., Blackwell P.G., Niu M., Buck C.E., Guilderson T.P., Heaton T.J., Palmer
J.G. et al. (2013) SHCal13 Southern Hemisphere calibration, 0–50,000 cal yr BP.
Radiocarbon. 55: 1889–1903. https://doi.org/10.2458/azu_js_rc.55.16783
Holmgren, K., Karlén, W., Lauritzen, S. E., Lee-Thorp, J. A., Partridge, T. C., Piketh, S.,
Repinski, P., Stevenson, C., Svanered, O., & Tyson, P. D. (1999). A 3000-year high-
resolution stalagmite-based record of palaeoclimate for northeastern South Africa.
Holocene, 9(3), 295–309. https://doi.org/10.1191/095968399672625464
Hoscilo, A., Balzter, H., Bartholomé, E., Boschetti, M., Brivio, P. A., Brink, A., Clerici, M., &
Pekel, J. F. (2015). A conceptual model for assessing rainfall and vegetation trends in sub-
Saharan Africa from satellite data. International Journal of Climatology, 35(12), 3582–
3592. https://doi.org/10.1002/joc.4231
Hua, Q., Barbetti, M., & Rakowski, A. (2013). Atmospheric Radiocarbon for the Period 1950–
2010. Radiocarbon, 55(4), 2059-2072. https://doi.org/10.2458/azu_js_rc.v55i2.16177
Huffman, T. N., & Woodborne, S. (2016). Archaeology, baobabs and drought: Cultural proxies
and environmental data from the Mapungubwe landscape, southern Africa. Holocene,
26(3), 464–470. https://doi.org/10.1177/0959683615609753
Huffman, T.N. (2004) The archaeology of the Nguni past. Southern African Humanities.
16(1929): 79–111. https://hdl.handle.net/10520/EJC84742
Ingram, J. C., & Dawson, T. P. (2005). Inter-annual analysis of deforestation hotspots in
Madagascar from high temporal resolution satellite observations. International Journal of
Remote Sensing, 26(7), 1447–1461. https://doi.org/10.1080/01431160412331291189
IPCC (2021): Climate Change (2021) The Physical Science Basis. Contribution of Working
Group I to the Sixth Assessment Report of the Intergovernmental Panel on Climate Change
[Masson-Delmotte, V., Zhai, P., Pirani, A., Connors, S. L., Péan, C., Berger, S. , Caud, N.,
Chen, Y., Goldfarb, L., Gomis, M. I. , Huang, M., Leitzell, K., Lonnoy, ., E., Matthews, J.
B. R., Maycock, T. K., Waterfield, T., Yelekçi, O., Yu, R. & Zhou, B. (eds.)]. Cambridge
University Press.
.CC-BY 4.0 International licensemade available under a
(which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is
The copyright holder for this preprintthis version posted August 20, 2025. ; https://doi.org/10.1101/2025.08.15.670475doi: bioRxiv preprint
Johnson, T.C., Barry, S.L., Chan, Y., Wilkinson, P. 2001. Decadal record of climate variability
spanning the past 700 yr in the Southern Tropics of East Africa. Geology 29 (1): 83–86.
https://doi.org/10.1130/0091
Jury, M. R. (2016). Summer climate of Madagascar and monsoon pulsing of its vortex.
Meteorology and Atmospheric Physics, 128(1), 117–129. https://doi.org/10.1007/s00703-
015-0401-5
Jury, M. R., & Huang, B. (2004). The Rossby wave as a key mechanism of Indian Ocean climate
variability. Deep Sea Research Part I: Oceanographic Research Papers, 51(12), 2123–2136.
https://doi.org/10.1016/j.dsr.2004.06.005
Kaufmann, J. C. & Tsirahamba, S. (2006). Forests and Thorns: Conditions of Change Affecting
Mahafale Pastoralists in Southwestern Madagascar. Conservation and Society. 4(2): 231–
261.
King, J., Anchukaitis, K.J., Allen, K. et al. Trends and variability in the Southern Annular Mode
over the Common Era. Nat Commun 14, 2324 (2023). https://doi.org/10.1038/s41467-023-
37643-1
Kuiper, M., Meijerink, G., & Eaton, D. (2007). Rural livelihoods: Interplay between farm
activities, non-farm activities and the resource base. Science for Agriculture and Rural
Development in Low-Income Countries, 77–95. https://doi.org/10.1007/978-1-4020-6617-
7_5
Lechleitner, F. A., Breitenbach, S. F. M., Rehfeld, K., Ridley, H. E., Asmerom, Y., Prufer, K.
M., Marwan, N., Goswami, B., Kennett, D. J., Aquino, V. V., Polyak, V., Haug, G. H.,
Eglinton, T. I., & Baldini, J. U. L. (2017). Tropical rainfall over the last two millennia:
Evidence for a low-latitude hydrologic seesaw. Scientific Reports, 7(June 2016), 1–9.
https://doi.org/10.1038/srep45809
Li, J., Xie, S.P., Cook, E. et al. (2011). Interdecadal modulation of El Niño amplitude during the
past millennium. Nature Climate Change. 1, 114–118.
https://doi.org/10.1038/nclimate1086
.CC-BY 4.0 International licensemade available under a
(which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is
The copyright holder for this preprintthis version posted August 20, 2025. ; https://doi.org/10.1101/2025.08.15.670475doi: bioRxiv preprint
Liu, Z., Otto-Bliesner, Z., Kutzbach, J., Li, L. C. (2003). Shields Coupled climate simulation of
the evolution of global monsoons in the Holocene. Journal of Climate, 16, 2472-2490.
https://doi.org/10.1175/1520-0442(2003)0162.0.CO;2
Loader, N. J., Robertson, I., Barker, A. C., Switsur, V. R., & Waterhouse, J. S. (1997). An
improved technique for the batch processing of small wholewood samples to a-cellulose.
Chemical Geology, 136, 313–317. https://doi.org/10.1016/S0009-2541(96)00133-7
MacDonald, G. M., & Case, R. A. (2005). Variations in the Pacific Decadal Oscillation over the
past millennium. Geophysical Research Letters, 32(8), 1–4.
https://doi.org/10.1029/2005GL022478
Macron, C., Pohl, B., Richard, Y., & Bessafi, M. (2014). How do tropical temperate troughs
form and develop over Southern Africa? Journal of Climate, 27(4), 1633–1647.
https://doi.org/10.1175/JCLI-D-13-00175.1
Mamalakis, A., Randerson, J.T., Yu, JY. et al. (2021). Zonally contrasting shifts of the tropical
rain belt in response to climate change. Nature Climate Change 11, 143–151.
https://doi.org/10.1038/s41558-020-00963-x
Manhique, A. J., Reason, C. J. C., Rydberg, L. & Fauchereau, N. (2011). ENSO and Indian sea
surface temperatures with tropical temperate troughs over Mozambique and the southwest
Indian Ocean. International Journal of Climatology. 31, 1–13.
https://doi.org/10.1002/joc.2050
Mann, M. E., Zhang, Z., Rutherford, S., Bradley, R. S., Hughes, M. K., Shindell, D., Ammann,
C., Faluvegi, G., & Ni, F. (2009). Global Signatures and Dynamical Origins of the Little
Ice Age and Medieval Climate Anomaly. Science, 326(5957), 1256–1260.
https://doi.org/10.1126/science.1177303
Mason, S. J. & Jury, M. R. (1997). ‘Climatic change and variability over Southern Africa: A
Reflection on Underlying Processes’, Progress in Physical Geoggraphy. 21, 23–50.
https://doi.org/10.1177/030913339702100103
.CC-BY 4.0 International licensemade available under a
(which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is
The copyright holder for this preprintthis version posted August 20, 2025. ; https://doi.org/10.1101/2025.08.15.670475doi: bioRxiv preprint
Matsumoto, K. & Burney, D. A. (1994). Late Holocene environments at Lake Mitsinjo,
northwestern Madagascar. Holocene, 4(1), 16–24.
https://doi.org/10.1177/095968369400400103
McCarroll, D., & Loader, N. J. (2004). Stable isotopes in tree rings. Quaternary Science
Reviews, 23(7–8), 771–801. https://doi.org/10.1016/j.quascirev.2003.06.017
Miller, C., Finch, J., Hill, T., Peterse, F., Humphries, M., Zabel, M., and Schefuß, E. (2019) Late
Quaternary climate variability at Mfabeni peatland, eastern South Africa, ClimAte of the
Past, 15,1153–1170,
https://doi.org/10.5194/cp-15-1153-2019
Nakamura, N., Kayanne, H., Iijima, H., McClanahan, T.R., Behera, S.K., Yamagata, T., (2009).
Mode shift in the Indian Ocean climate under global warming stress. Geophysical Research
Letters. 36 http://dx.doi.org/10.1029/2009GL040590
Nash. (2017). Changes in Precipitation Over Southern Africa During Recent Centuries. Climate
Science. https://doi.org/10.1093/acrefore/9780190228620.013.539
Nasri, M., & Modarres, R. (2009). Dry spell trend analysis of Isfahan Province, Iran.
International Journal of Climatology, 29, 1430–1438. https://doi.org/10.1002/joc
Neukom, R., & Gergis, J. (2012). Southern Hemisphere high-resolution palaeoclimate records of
the last 2000 years. The Holocene, 22(5), 501–524.
https://doi.org/10.1177/0959683611427335
Nicholson, S. E., Klotter, D., & Dezfuli, A. K. (2012). Spatial reconstruction of semi-quantitative
precipitation fields over Africa during the nineteenth century from documentary evidence
and gauge data. Quaternary Research, 78(1), 13–23.
https://doi.org/10.1016/j.yqres.2012.03.012
PAGES 2k Consortium. 2013. Continental-scale temperature variability during the past two
millennia. Nature Geoscience. 6: 339-346. http://dx.doi.org/10.1038/1849t
Patrut, A., Reden, K. F. Von, Danthu, P., Pock-tsy, J. L., Rakosy, L., Patrut, R. T., Lowy, D. A.,
& Margineanu, D. (2015). Nuclear Instruments and Methods in Physics Research B AMS
radiocarbon dating of very large Grandidier ’ s baobabs ( Adansonia grandidieri ). Nuclear
.CC-BY 4.0 International licensemade available under a
(which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is
The copyright holder for this preprintthis version posted August 20, 2025. ; https://doi.org/10.1101/2025.08.15.670475doi: bioRxiv preprint
Inst. and Methods in Physics Research, B, 361, 591–598.
https://doi.org/10.1016/j.nimb.2015.04.044
Onyeaka, H., Nwauzoma, U. M., Akinsemolu, A. A., Tamasiga, P., Duan, K., Al-Sharify, Z. T.,
& Siyanbola, K. F. (2024). The ripple effects of climate change on agricultural
sustainability and food security in Africa. Food and Energy Security, 13, e567.
https://doi.org/10.1002/fes3.567
Patrut, A., Woodborne, S., Von Reden, K. F., Hall, G., Patrut, R. T., Rakosy, L., Danthu, P.,
Pock-Tsy, J. M. L., Lowy, D. A., & Margineanu, D. (2017). The growth stop phenomenon
of baobabs (Adansonia spp.) Identified by radiocarbon dating. Radiocarbon, 59(2), 435–
448.
https://doi.org/10.1017/RDC.2016.92
Pohlert, T. (2018). R Package “trends.” 18. https://cran.r-
project.org/web/packages/trend/vignettes/trend.pdf
Putnam, A. E., & Broecker, W. S. (2017). Human-induced changes in the distribution of rainfall.
Science Advances, 3(5), 1–14. https://doi.org/10.1126/sciadv.1600871
Railsback, L. B., Brook, G. A., Liang, F., Voarintsoa, N. R. G., Cheng, H., & Edwards, R. L.
(2018). A multi-proxy climate record from a northwestern Botswana stalagmite suggesting
wetness late in the Little Ice Age (1810–1820 CE) and drying thereafter in response to
changing migration of the tropical rain belt or ITCZ. Palaeogeography, Palaeoclimatology,
Palaeoecology, 506(April), 139–153. https://doi.org/10.1016/j.palaeo.2018.06.029
Ramanantsoa, J. D., Penven, P.,Krug, M., Gula, J., & Rouault, M. (2018). Uncovering a new
current: The Southwest Madagascar Coastal Current. Geophysical Research Letters, 45,
1930–1938.
https://doi.org/10.1002/2017GL075900
Ratna, S. B., Behera, S., Ratnam, J. V. , Takahasgi, K., & Yamagata, T. (2012) An index for
tropical temperate troughs over southern Africa. Climate Dynamics, 41, 421–441.
http://dx.doi.org/10.1007/s00382-012-1540-8 .
Razanamaro O., Rasoamanana E., Rakouth B., Randriamalala J.R., Rabakonadrianina E.,
Clément-Vidal A., Leong Pock Tsy J.M., Menut C., Danthu P. (2015). Chemical
.CC-BY 4.0 International licensemade available under a
(which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is
The copyright holder for this preprintthis version posted August 20, 2025. ; https://doi.org/10.1101/2025.08.15.670475doi: bioRxiv preprint
characterization of floral scents in the six endemic baobab species (Adansonia sp.) of
Madagascar. Biochemical Systematics and Ecology, 60 : 238-248
http://dx.doi.org/10.1016/j.bse.2015.04.005
Razanatsoa, E., Gillson,L., Virah-Sawmy, M., and Woodborne, S. (2021) Pollen records of the
14th and 20th centuries AD from Lake Tsizavatsy in southwest Madagascar.
Palaeoecology of Africa, 35, pp. 309-315. http://dx.doi.org/10.1201/9781003162766-20
Razanatsoa, E., Gillson, L., Virah-Sawmy, M., and Woodborne, S. (2022) Synergy between
climate and human land-use maintained open vegetation in southwest Madagascar over the
last millennium. The Holocene. 31(12). https://doi.org/10.1177/09596836211041731
Reason, C. J. C., & Mulenga, H. (1999). Relationships between South African rainfall and SST
anomalies in the southwest Indian Ocean. International Journal of Climatology, 19(15),
1651–1673.
https://doi.org/10.1002/(SICI)1097-0088(199912)19:153.0.CO;2-U
Robertson, I., Loader, N., Froyd, C., Zambatis, N., Whyte, I., & S, W. (2006). The potential of
the baobab (Adansonia digitata L.) as a proxy climate archive. Applied Geochemistry, 21,
1674–1680. https://doi.org/10.1016/j.apgeochem.2006.07.005
Russell, J. M. & Johnson, T. C. (2007). Little ice age drought in equatorial Africa: inter- tropical
convergence zone migrations and El Niño southern oscillation variability. Geology. 35: 21-
24. http://dx.doi.org/10.1130/G23125A.1.
Russell, J. M., Verschuren, D., & Eggermont, H. (2007). Spatial complexity of “Little Ice Age”
climate in East Africa: Sedimentary records from two crater lake basins in western
Uganda. Holocene, 17(2), 183–193. https://doi.org/10.1177/0959683607075832
Sachs, J. P., Sachse, D., Smittenberg, R. H., Zhang, Z., Battisti, D. S., & Golubic, S. (2009).
Southward movement of the Pacific intertropical convergence zone AD 1400-1850. Nature
Geoscience, 2(7), 519–525. https://doi.org/10.1038/ngeo554
Saji, N. H., & Yamagata, T. (2003). Possible impacts of Indian Ocean Dipole mode events on
global climate. Climate Research, 25(2), 151–169. https://doi.org/10.3354/cr025151
.CC-BY 4.0 International licensemade available under a
(which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is
The copyright holder for this preprintthis version posted August 20, 2025. ; https://doi.org/10.1101/2025.08.15.670475doi: bioRxiv preprint
Schneider, T., Bischoff, T. & Haug, G. (2014). Migrations and dynamics of the intertropical
convergence zone. Nature, 513, 45–53. https://doi.org/10.1038/nature13636
Scroxton, N., Burns, S. J., Mcgee, D., Hardt, B., Godfrey, L. R., Ranivoharimanana, L., & Faina,
P. (2017). Hemispherically in-phase precipitation variability over the last 1700 years in a
Madagascar speleothem record. 164. https://doi.org/10.1016/j.quascirev.2017.03.017
Serele, C., Pérez-Hoyos, A., & Kayitakire, F. (2020). Mapping of groundwater potential zones in
the drought-prone areas of south Madagascar using geospatial techniques. Geoscience
Frontiers, 11(4), 1403–1413. https://doi.org/10.1016/j.gsf.2019.11.012
Sigman, D. M., Hain, M. P., and Haug, G. H. (2010). The polar ocean and glacial cycles in
atmospheric CO2 concentration, Nature, 466, 47–55, https://doi.org/10.1038/nature09149,
201
Stager, J. C., Ryves, D., Cumming, B. F., David Meeker, L., & Beer, J. (2005). Solar variability
and the levels of Lake Victoria, East Africa, during the last millenium. Journal of
Paleolimnology, 33(2), 243–251. https://doi.org/10.1007/s10933-004-4227-2
Svensson, A., Andersen, K. K., Bigler, M., Clausen, H. B., Dahl-Jensen, D., Davies, S. M.,
Johnsen, S. J., Muscheler, R., Rasmussen, S. O., Röthlisberger, R., Peder Steffensen, J., &
Vinther, B. M. (2006). The Greenland Ice Core Chronology 2005, 15-42 ka. Part 2:
comparison to other records. Quaternary Science Reviews, 25(23–24), 3258–3267.
https://doi.org/10.1016/j.quascirev.2006.08.003
Tadross, M., Randriamarolaza, L., Rabefitia, Z., & Ki Yip, Z. (2008). Climate change in
Madagascar; recent past and future. … DC (World Bank), February, 18.
http://www.mediagrapher.org/gripweb/sites/default/files/disaster_risk_profiles/Madagascar
Climate Report.pdf
Tambo, J.A. & Abdoulaye, T. (2013). Smallholder farmers’ perceptions of and adaptations to
climate change in the Nigerian savanna. Regional Environmental Change 13, 375–388
https://doi.org/10.1007/s10113-012-0351-0
.CC-BY 4.0 International licensemade available under a
(which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is
The copyright holder for this preprintthis version posted August 20, 2025. ; https://doi.org/10.1101/2025.08.15.670475doi: bioRxiv preprint
Taylor, A. K., Berke, M. A., Castañeda, I. S., Koutsodendris, A., Campos, H., Hall, I. R.,
Hemming, S. R., LeVay, L. J., Sierra, A. C., & O’Connor, K. (2021). Plio‐Pleistocene
Continental Hydroclimate and Indian Ocean Sea Surface Temperatures at the Southeast
African Margin. Paleoceanography and Paleoclimatology, 36(3), 1–18.
https://doi.org/10.1029/2020pa004186
Thomas, D.S.G., Twyman, C., Osbahr, H. et al. (2007). Adaptation to climate change and
variability: farmer responses to intra-seasonal precipitation trends in South Africa. Climatic
Change 83, 301–322 https://doi.org/10.1007/s10584-006-9205-4
Thompson, D. W. J., & Wallace, J. M. (2000). Annular modes in the extratropical circulation.
Part I: Month-to-month variability. Journal of Climate, 13(5), 1000–1016.
https://doi.org/10.1175/1520-0442(2000)0132.0.CO;2
Thompson, L. G., Mosley-Thompson, E., Davis, M. E., Lin, P. N., Henderson, K., & Mashiotta,
T. A. (2003). Tropical glacier and ice core evidence of climate change on annual to
millennial time scales. Climatic Change, 59(1–2), 137–155.
https://doi.org/10.1023/A:1024472313775
Tierney, J. E., Smerdon, J. E., Anchukaitis, K. J. & Seager, R. (2013). Multidecadal variability in
East African hydroclimate controlled by the Indian Ocean. Nature. 493: 389-392.
http://dx.doi.org/10.1038/nature11785
Tieszen, L. L. (1991). Natural variations in the carbon isotope values of plants: implications for
archaeology, ecology and palaeoecology. Journal of Archaeological Science. 18: 227–248.
https://doi.org/10.1016/0305-4403(91)90063-U
Trenberth, K.E. (1979). Interannual variability of the 500 mb zonal-mean flow in the Southern
Hemisphere. Monthly Weather Reviews. 107, 1515–1524. https://doi.org/10.1175/1520-
0493(1979)1072.0.CO;2
Tsen, E.W.J., Sitzia, T. and Webber, B.L. (2016). To core, or not to core: the impact of coring on
tree health and a best-practice framework for collecting dendrochronological information
from living trees. Biological Review, 91: 899-924. https://doi.org/10.1111/brv.12200
.CC-BY 4.0 International licensemade available under a
(which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is
The copyright holder for this preprintthis version posted August 20, 2025. ; https://doi.org/10.1101/2025.08.15.670475doi: bioRxiv preprint
Tyson, P. D., Odada, E. O., & Partridge, T. C. (2001). Late quaternary environmental change in
southern Africa. South African Journal of Science, 97(3–4), 139–150.
https://hdl.handle.net/10520/EJC97298
Verschuren D. (2003). Lake-based climate reconstruction in Africa: progress and challenges. In:
Martens K. (eds) Aquatic Biodiversity. Developments in Hydrobiology, vol 171. Springer,
Dordrecht. https://doi.org/10.1007/978-94-007-1084-9_22
Verschuren, D., Laird, K. R., & Cumming, B. F. (2000). Rainfallanddroughtinequatorialeast
Africa during the past 1, 100 years. Solar Cells, 403(January).
https://doi.org/10.1038/35000179
Vigaud, N., Richard, Y., Rouault, M. & Fauchereau, N. (2007). Water vapour transport from the
tropical Atlantic and summer rainfall in tropical southern Africa. Climate Dynamics, 28,
113–123. https://doi.org/10.1007/s00382-006-0186-9
Virah-Sawmy, M., Gillson, L., Gardner, C. J., Anderson, A., Clark, G., & Haberle, S. (2016). A
landscape vulnerability framework for identifying integrated conservation and adaptation
pathways to climate change: the case of Madagascar’s spiny forest. Landscape Ecology,
31(3), 637–654. https://doi.org/10.1007/s10980-015-0269-2
Voarintsoa, N. R. G., Wang, L., Railsback, L. B., Brook, G. A., Liang, F., Cheng, H., &
Edwards, R. L. (2017). Multiple proxy analyses of a U/Th-dated stalagmite to reconstruct
paleoenvironmental changes in northwestern Madagascar between 370 CE and 1300 CE.
Palaeogeography, Palaeoclimatology, Palaeoecology, 469, 138–155.
https://doi.org/10.1016/j.palaeo.2017.01.003
Von Heland, J., & Folke, C. (2014). A social contract with the ancestors-Culture and ecosystem
services in southern Madagascar. Global Environmental Change, 24(1), 251–264.
https://doi.org/10.1016/j.gloenvcha.2013.11.003
Wang, G., & Feng, X. (2012). Response of plants’ water use efficiency to increasing atmospheric
CO2 concentration. Environmental Science & Technology, 46(16), 8610–8620.
https://doi.org/10.1021/es301323m
.CC-BY 4.0 International licensemade available under a
(which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is
The copyright holder for this preprintthis version posted August 20, 2025. ; https://doi.org/10.1101/2025.08.15.670475doi: bioRxiv preprint
Wang, L., Brook, G. A., Burney, D. A., Voarintsoa, N. R. G., Liang, F., Cheng, H., & Edwards,
R. L. (2019). The African Humid Period, rapid climate change events, the timing of human
colonization, and megafaunal extinctions in Madagascar during the Holocene: Evidence
from a 2m Anjohibe Cave stalagmite. Quaternary Science Reviews, 210, 136–153.
https://doi.org/10.1016/j.quascirev.2019.02.004
Watanabe, T. K., Watanabe, T., Yamazaki, A., Pfeiffer, M., & Claereboudt, M. R. (2019). Oman
coral δ 18 O seawater record suggests that Western Indian Ocean upwelling uncouples
from the Indian Ocean Dipole during the global-warming hiatus. Scientific Reports, 9(1),
1–9.
https://doi.org/10.1038/s41598-018-38429-y
Weathering Risk (W.R). (2023). Climate Risk Profile for Southern Africa.
Wils THG, Robertson I, Woodborne S, Hall G, Koprowski M, Eshetu Z. 2016. Anthropogenic
forcing increases the water-use efficiency of African trees. Journal of Quaternary Science
31, 386-390. https://doi.org/10.1002/jqs.2865
Woodborne S, Gandiwa P, Hall G, Patrut A, Finch J (2016) A Regional Stable Carbon Isotope
Dendro-Climatology from the South African Summer Rainfall Area. PLoS ONE 11(7):
e0159361. https://doi.org/10.1371/journal.pone.0159361
Woodborne, S., Hall, G., Robertson, I., Patrut, A., Rouault, M., Loader, N. J., & Hofmeyr, M.
(2015). A 1000-year carbon isotope rainfall proxy record from South African baobab trees
(Adansonia digitata L.). PLoS ONE, 10(5), 1–18.
https://doi.org/10.1371/journal.pone.0124202
World Bank Group (2017). World Bank Annual Report (English). Washington, D.C. :.
http://documents.worldbank.org/curated/en/143021506909711004/World-Bank-Annual-
Report-2017
Yaro, J. A., Teye, J., & Bawakyillenuo, S. (2014). Local institutions and adaptive capacity to
climate change/variability in the northern savannah of Ghana. Climate and Development,
7(3), 235–245.
https://doi.org/10.1080/17565529.2014.951018
.CC-BY 4.0 International licensemade available under a
(which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is
The copyright holder for this preprintthis version posted August 20, 2025. ; https://doi.org/10.1101/2025.08.15.670475doi: bioRxiv preprint
Zinke, J., Loveday, B. R., Reason, C. J. C., Kroon, D., Ocean, S. I., & Age, L. I. (2014).
temperature variability in the Agulhas Current core region over the past 334. Scientific
reports. pp. 1–8. https://doi.org/10.1038/srep04393
4. FUNDING
This project has been funded as part of the Faculty PhD fellowship (University of Cape Town,
R.E.) 2015-2018 and the Applied Centre for Climate and Earth Systems Science (ACCESS NRF
UID 98018, R.E.) project, the UCT University Research Committee accredited (URC) and
COVID supplemental support from the University of Cape Town [URC, 2019-2020] and the
NRF/SASSCAL (Southern African Science Service Centre for Climate Change and Adaptive
Land Management, grant number 118589), the NRF/African Origins Platform (grant number
117666), and NRF Competitive Programme for Rated Researchers (Grant Number 118538).
Supplementary Information
SI1: Radiocarbon dates of each tree replicate from the four trees collected in southwest
Madagascar
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