Wildlife uses and hunting patterns in rural communities of the Yucatan Peninsula, Mexico

Background

Use of plant resources and ecosystems practiced by indigenous peoples of Mesoamerica commonly involves domestication of plant populations and landscapes. Our study analyzed interactions of coexisting wild and managed populations of the pitaya Stenocereus pruinosus, a columnar cactus used for its edible fruit occurring in natural forests, silviculturally managed in milpa agroforestry systems, and agriculturally managed in homegardens of the Tehuacán Valley, Mexico. We aimed at analyzing criteria of artificial selection and their consequences on phenotypic diversity and differentiation, as well as documenting management of propagules at landscape level and their possible contribution to gene flow among populations.

Methods

Semi-structured interviews were conducted to 83 households of the region to document perception of variation, criteria of artificial selection, and patterns of moving propagules among wild and managed populations. Morphological variation of trees from nine wild, silviculturally and agriculturally managed populations was analyzed for 37 characters through univariate and multivariate statistical methods. In addition, indexes of morphological diversity (MD) per population and phenotypic differentiation (PD) among populations were calculated using character states and frequencies.

Results

People recognized 15 pitaya varieties based on their pulp color, fruit size, form, flavor, and thorniness. On average, in wild populations we recorded one variety per population, in silviculturally managed populations 1.58 ± 0.77 varieties per parcel, and in agriculturally managed populations 2.19 ± 1.12 varieties per homegarden. Farmers select in favor of sweet flavor (71% of households interviewed) and pulp color (46%) mainly red, orange and yellow. Artificial selection is practiced in homegardens and 65% of people interviewed also do it in agroforestry systems. People obtain fruit and branches from different population types and move propagules from one another. Multivariate analyses showed morphological differentiation of wild and agriculturally managed populations, mainly due to differences in reproductive characters; however, the phenotypic differentiation indexes were relatively low among all populations studied. Morphological diversity of S. pruinosus (average MD = 0.600) is higher than in other columnar cacti species previously analyzed.

Conclusions

Artificial selection in favor of high quality fruit promotes morphological variation and divergence because of the continual replacement of plant material propagated and introduction of propagules from other villages and regions. This process is counteracted by high gene flow influenced by natural factors (pollinators and seed dispersers) but also by human management (movement of propagules among populations), all of which determines relatively low phenotypic differentiation among populations. Conservation of genetic resources of S. pruinosus should be based on the traditional forms of germplasm management by local people.     

Evolution under domestication: ongoing artificial selection and divergence of wild and managed Stenocereus pruinosus (Cactaceae) populations in the Tehuac��n Valley, Mexico

Background and Aims

The Tehuacán Valley in Mexico is a principal area of plant domestication in Mesoamerica. There, artificial selection is currently practised on nearly 120 native plant species with coexisting wild, silvicultural and cultivated populations, providing an excellent setting for studying ongoing mechanisms of evolution under domestication. One of these species is the columnar cactus Stenocereus pruinosus, in which we studied how artificial selection is operating through traditional management and whether it has determined morphological and genetic divergence between wild and managed populations.

Methods

Semi-structured interviews were conducted with 83 households of three villages to investigate motives and mechanisms of artificial selection. Management effects were studied by comparing variation patterns of 14 morphological characters and population genetics (four microsatellite loci) of 264 plants from nine wild, silvicultural and cultivated populations.

Key Results

Variation in fruit characters was recognized by most people, and was the principal target of artificial selection directed to favour larger and sweeter fruits with thinner or thicker peel, fewer spines and pulp colours others than red. Artificial selection operates in agroforestry systems favouring abundance (through not felling plants and planting branches) of the preferred phenotypes, and acts more intensely in household gardens. Significant morphological divergence between wild and managed populations was observed in fruit characters and plant vigour. On average, genetic diversity in silvicultural populations (H_E = 0·743) was higher than in wild (H_E = 0·726) and cultivated (H_E = 0·700) populations. Most of the genetic variation (90·58 %) occurred within populations. High gene flow (Nm_FST > 2) was identified among almost all populations studied, but was slightly limited by mountains among wild populations, and by artificial selection among wild and managed populations.

Conclusions

Traditional management of S. pruinosus involves artificial selection, which, despite the high levels of gene flow, has promoted morphological divergence and moderate genetic structure between wild and managed populations, while conserving genetic diversity.     

Traditional Management and Morphological Patterns of <Emphasis Type="Italic">Myrtillocactus schenckii</Emphasis> (Cactaceae) in the Tehuacán Valley,Central Mexico<Superscript>1</Superscript>

Traditional Management and Morphological Patterns of Myrtillocactus schenckii (Cactaceae) in the Tehuacán Valley, Central Mexico Myrtillocactus schenckii is a columnar cactus endemic to central México and appreciated for its edible fruit. In the Tehuacán Valley it occurs wild in thorn-scrub forests, but it is also under silviculture management—tolerate in agroforestry systems and cultivated in home gardens. Ethnobotanical and morphometric studies were conducted to document its use and management forms, as well as consequences of artificial selection on morphological patterns in managed populations. In silviculture populations artificial selection occurs through selective tolerance of the phenotypes producing more, larger, and sweeter fruits, whereas in home gardens people plant branches of the preferred phenotypes. Morphological differences were found especially in fruit size and production, the main targets of artificial selection. The highest average values of fruit size and production were found in cultivated populations (0.993 cm³ and 204.45 fruits per branch, respectively), intermediate in silviculture populations (0.819 cm³ and 70.63 fruits per branch), and the lowest in the wild (0.68 cm³ and 59.75 fruits per branch). Multivariate statistical analyses differentiated populations according to their management type. Morphological diversity was higher in managed populations (0.703 ± 0.029 and 0.679 ± 0.019 in silviculture and cultivated populations, respectively) than in the wild (0.652 ± 0.016). Managed plants of M. schenckii do not show signs of depending on humans for survival and reproduction and, therefore, domestication should be considered incipient.     

Morphological variation and the process of domestication of Stenocereus stellatus (Cactaceae) in Central Mexico

Morphological variation was analyzed in wild, managed in situ, and cultivated populations of the columnar cactus Stenocereus stellatus in central Mexico. The purpose was to evaluate whether morphological divergence between manipulated and wild populations has resulted from domestication processes. Variation of 23 morphological characters was analyzed among 324 individuals from 19 populations of the Tehuacán Valley and La Mixteca Baja. Multivariate statistical analyses were used to group individuals and populations according to their morphological similarity. Individuals grouped according to the way of management and fruit characteristics were the most relevant for grouping. Within each region, sweet fruits with pulp colors other than red were more frequent in cultivated populations, where fruits were also larger, contained more and bigger seeds, and had thinner peel and fewer spines than fruits from wild individuals. Phenotypes common in managed in situ and cultivated populations generally occur in the wild but in lower frequencies. Artificial selection has thus operated by enhancing and maintaining desirable rare phenotypes in managed in situ and cultivated populations, causing divergent patterns of morphological variation from wild populations. Cultivation has caused the strongest level of divergence, but divergence has also been significant with management of wild populations in situ.     

Maintenance of Phenotypic and Genotypic Diversity in Managed Populations of Stenocereus Stellatus (Cactaceae) by Indigenous Peoples in Central Mexico

The columnar cactus Stenocereus stellatus is used in Central Mexico for its edible fruits which are harvested in wild, managed in situ and cultivated populations. Management in situ of wild populations is conducted by selectively sparing and enhancing the abundance of plants with desirable phenotypes when fields are cleared for agricultural use. Cultivation of desirable phenotypes is carried out by vegetative propagation in homegardens. Effects of human management on morphological and genetic variation of S. stellatus were analyzed by comparing morphological diversity indices (MD, based on Simpson’s index) and expected (H_e) heterozygosity indices from allozyme analysis, in wild, managed in situ, and cultivated populations from La Mixteca and the Tehuacán Valley regions. Morphological diversity was similar among regions, but populations from the wetter La Mixteca region averaged higher genetic variation (H_e = 0.279) than populations from Tehuacán (H_e = 0.265). On average, populations manipulated by people had higher levels of variation (MD = 0.479 ± 0.012, H_e = 0.289 in cultivated populations; MD = 0.461 ± 0.014, H_e = 0.270 in managed in situ populations) than wild populations (MD = 0.408 ± 0.017, H_e = 0.253), which is apparently due to a continual introduction and replacement of plant materials in the manipulated populations. The results illustrate that human management may not only maintain but also increase both morphological and genetic diversity of manipulated plant populations in relation to that existing in the wild. Managed in situ and cultivated populations of S. stellatus are important reservoirs of variation, and are crucial for the general maintenance of diversity in wild populations. These populations may play a principal role in designing strategies for the conservation of variation of this cactus.     

Round and large: morphological and genetic consequences of artificial selection on the gourd tree Crescentia cujete by the Maya of the Yucatan Peninsula, Mexico

Background and Aims

Artificial selection, the main driving force of domestication, depends on human perception of intraspecific variation and operates through management practices that drive morphological and genetic divergences with respect to wild populations. This study analysed the recognition of varieties of Crescentia cujete by Maya people in relation to preferred plant characters and documents ongoing processes of artificial selection influencing differential chloroplast DNA haplotype distribution in sympatric wild and home-garden populations.

Methods

Fifty-three home gardens in seven villages (93 trees) and two putative wild populations (43 trees) were sampled. Through semi-structured interviews we documented the nomenclature of varieties, their distinctive characters, provenance, frequency and management. Phenotypic divergence of fruits was assessed with morphometric analyses. Genetic analyses were performed through five cpDNA microsatellites.

Key Results

The Maya recognize two generic (wild/domesticated) and two specific domesticated (white/green) varieties of Crescentia cujete. In home gardens, most trees (68 %) were from domesticated varieties while some wild individuals (32 %) were tolerated. Cultivation involves mainly vegetative propagation (76 %). Domesticated fruits were significantly rounder, larger and with thicker pericarp than wild fruits. Haplotype A was dominant in home gardens (76 %) but absent in wild populations. Haplotypes B–F were found common in the wild but at low frequency (24 %) in home gardens.

Conclusions

The gourd tree is managed through clonal and sexual propagules, fruit form and size being the main targets of artificial selection. Domesticated varieties belong to a lineage preserved by vegetative propagation but propagation by seeds and tolerance of spontaneous trees favour gene flow from wild populations. Five mutational steps between haplotypes A and D suggest that domesticated germplasm has been introduced to the region. The close relationship between Maya nomenclature and artificial selection has maintained the morphological and haplotypic identity (probably for centuries) of domesticated Crescentia despite gene flow from wild populations.     

Phenotypic differentiation between wild and domesticated varieties of <Emphasis Type="Italic">Crescentia cujete</Emphasis>L. and culturally relevant uses of their fruits as bowls in the Yucatan Peninsula,Mexico

Background

Selection criteria are important for analyzing domestication of perennial plant species, which experience a selection pressure throughout several human generations. We analyze the preferred morphological characteristics of Crescentia cujete fruits, which are used as bowls by the Maya of Yucatan, according to the uses they are given and the phenotypic consequences of artificial selection between one wild and three domesticated varieties.

Methods

We performed 40 semi-structured interviews in seven communities. We calculated Sutrop’s salience index (S) of five classes of ceremonial and daily life uses, and of each item from the two most salient classes. We sampled 238 bowls at homes of people interviewed and compared their shape, volume and thickness with 139 fruits collected in homegardens and 179 from the wild. Morphology of varieties was assessed in fruit (n?=?114 trees) and vegetative characters (n?=?136 trees). Differences between varieties were evaluated through linear discriminant analysis (LDA).

Results

Use of bowls as containers for the Day of the Dead offerings was the most salient class (S?=?0.489) with chocolate as its most salient beverage (S?=?0.491), followed by consumption of daily beverages (S?=?0.423), especially maize-based pozol (S?=?0.412). The sacred saka’ and balche' are offered in different sized bowls during agricultural and domestic rituals. Roundness was the most relevant character for these uses, as bowls from households showed a strong selection towards round shapes compared with wild and homegarden fruits. Larger fruits from domesticated varieties were also preferred over small wild fruits, although in the household different sizes of the domesticated varieties are useful. LDA separated wild from domesticated trees (p?<?0.001) according to both fruit and vegetative variables, but domesticated varieties were not different among themselves.

Conclusions

The association between C. cujete bowls and traditional beverages in ritual and daily life situations has driven for centuries the selection of preferred fruit morphology in this tree. Selection of fruit roundness and volume has allowed for the differentiation between the wild variety and the three domesticated ones, counteracting gene flow among them. By choosing the best fruits from domesticated varieties propagated in homegardens, the Maya people model the domestication process of this important tree in their culture.

     

Evolution of Polaskia chichipe (Cactaceae) under domestication in the Tehuacan Valley, central Mexico: reproductive biology

Polaskia chichipe, a columnar cactus, is cultivated for its edible fruits in central Mexico. This study analyzed whether artificial selection has modified its reproduction patterns and caused barriers to pollen exchange between wild, managed in situ, and cultivated populations. Anthesis was diurnal (～16 h in winter, ～10 h in spring) as well as partly nocturnal (～12 h in winter, ～3 h in spring), and flowers were pollinated by bees, hummingbirds, and hawk moths. Manual cross-pollination was ～37-49% effective in all populations. Self-pollination was ～12% successful in the wild, but twice as successful (～22-27%) in managed and cultivated populations. Diurnal pollination was ～35-55% effective in winter and 100% in spring. Nocturnal pollination was successful only in winter (15%). Crosses among individuals were more effective within populations than among populations, including populations under a similar management regimen. The least successful crosses were between wild and cultivated populations. Flowers were produced in all populations from January to March, but flowering peaks differed by 1 mo among wild, managed, and cultivated populations and by 2 mo between wild and cultivated populations. The latter interrupted pollen exchange in May. Seeds from managed and cultivated populations germinated faster than those from wild individuals. Domestication has seemingly favored self-compatible P. chichipe plants with higher fruit yield, a longer period of fruit production, and faster seed germination, attributes that have resulted in partial reproductive barriers between wild and manipulated populations.     

重庆地区野生大豆资源区域群居表型多样性初步分析

调查了重庆地区若干野生大豆区域居群的8个形态性状,分析了区域居群的表型多样性。结果显示,数量性状有比较丰富的多样性;区域居群的形态多样性存在一定的地域性(地理相关性);数量性状变异系数大小顺序依次为:百粒重>单株产量>地上部分茎叶干重>株高>播种至开花天数;各形态多样性指数(香农指数)显示,东北部区域>中部区域>西部区域的趋势。通过聚类分析,样本数较多的5个区域居群分为3个地理组和1个特别居群组,显示重庆地区地理区域居群存在明显的西部、中部、东北部生态地理的分化,东北部是形态变异丰富的地区。     

基于种实性状的无患子天然群体表型多样性研究

无患子(Sapindus mukorossi Gaertn.)是我国长江以南地区传统的重要绿化树种,其果皮富含皂苷,种仁富含油脂,是国家林业局审定的新型木本油料树种之一。为揭示无患子群体间和群体内种实表型性状变异式样,采用单因素方差分析、巢式方差分析、相关分析和主成分分析等方法,对无患子13个天然群体的10个种实表型性状进行比较分析,研究其群体间和群体内种实表型多样性以及表型变异与地理生态因子间的相关性。结果表明:无患子种实表型性状变异系数平均为7.34%,在群体间和群体内均存在丰富的表型变异。表型性状分化系数平均为62.21%,群体间变异(39.93%)大于群体内(27.46%),是无患子种实表型性状变异的主要来源。多数性状在群体间差异显著,不同程度的表现出边缘群体易于分化的特点,但地理变化规律不连续,而在群体内不同性状的差异性亦不一致。种子形态受群体地理生态的影响较果实形态大,西北部群体种子趋于椭圆形,东部、南部则趋于圆球形;地理纬度、年平均气温与多数种实性状间呈显著相关,是无患子种实表型性状变异的主要地理生态影响因子。可见,无患子种实表型性状在群体间、群体内变异都较为丰富,这些变异是由遗传和环境因素共同作用的结果。群体间和群体内多层次的变异为无患子优良种质资源保育和利用提供了物质基础。     

Contrasting patterns of body shape and neutral genetic divergence in marine and lake populations of threespine sticklebacks

Comparisons of neutral marker and quantitative trait divergence can provide important insights into the relative roles of natural selection and neutral genetic drift in population differentiation. We investigated phenotypic and genetic differentiation among Fennoscandian threespine stickleback (Gasterosteus aculeatus) populations, and found that the highest degree of differentiation occurred between sea and freshwater habitats. Within habitats, morphological divergence was highest among the different freshwater populations. Pairwise phenotypic and neutral genetic distances among populations were positively correlated, suggesting that genetic drift may have contributed to the morphological differentiation among habitats. On the other hand, the degree of phenotypic differentiation (PST) clearly surpassed the neutral expectation set by FST, suggesting a predominant role for natural selection over genetic drift as an explanation for the observed differentiation. However, separate PST/FST comparisons by habitats revealed that body shape divergence between lake and marine populations, and even among marine populations, can be strongly influenced by natural selection. On the other hand, genetic drift can play an important role in the differentiation among lake populations.     

Effects of natural and artificial selection on survival of columnar cacti seedlings: the role of adaptation to xeric and mesic environments

Escontria chiotilla, Polaskia chichipe, and Stenocereus pruinosus are species of Mexican columnar cacti that are economically important because of their edible fruits. These species are managed by gathering fruits from the wild, silvicultural management in agroforestry systems, and cultivation in home gardens. Previous studies reported that artificial selection favored individuals that produced larger fruits, which indirectly led to the production of larger seeds and seedlings, with possible effects on survival. We hypothesized that seedlings from managed populations would be larger but more susceptible to xeric conditions than those from wild populations. We evaluated the effects of artificial and natural selection on seedling survival of the three species in wild and managed populations, which were managed with low and high intensity, respectively. We tested seedling performance in gradients of shade (0, 40, and 80%) and humidity (low and high). A GLM of seedling survival showed significant differences among species, shade, and humidity treatments, with each species having environmental requirements associated with their particular adaptations. High humidity decreased seedling survival of all species, and high solar radiation decreased survival of S. pruinosus and P. chichipe. The effect of management type was significant only in S. pruinosus. Significant differences in the initial growth of seedlings among species were detected with ANOVA. In optimal conditions, the hypocotyl and the cotyledons decreased in size and the epicotyl grew, whereas under stress, these structures remained unchanged. The optimum conditions of shade and humidity varied among species and management types. The seedlings of S. pruinosus were the largest and the most susceptible, but in all species, seedlings from managed populations were more susceptible to environmental conditions. Thus, artificial selection influenced the susceptibility of these cacti to xeric environments.     

Population structure and genetic diversity in tristylous Narcissus triandrus: insights from microsatellite and chloroplast DNA variation

We investigated cpDNA sequence and nuclear microsatellite variation among populations of the wild daffodil Narcissus triandrus to examine the role of historical vs. contemporary forces in shaping population structure, morphological differentiation and sexual-system evolution. This wide-ranging heterostylous species of the Iberian Peninsula is largely composed of two allopatric varieties (vars. cernuus and triandrus), and populations with either stylar trimorphism or dimorphism. Dimorphic populations only occur in var. triandrus, are mainly restricted to the northwestern portion of the species range, and uniformly lack the mid-styled morph (M-morph). Chloroplast DNA (cpDNA) sequence variation revealed strong geographical structuring and evidence for a fragmentation event associated with differentiation of the two varieties. In var. triandrus, population fragmentation, restricted gene flow and isolation-by-distance were also inferred. Significant differences in genetic diversity and population structure between the two varieties likely reflect historical and contemporary differences in demography and gene flow among populations. Discordance between cpDNA markers and both microsatellite and morphological variation indicate that hybridization has occurred between the two varieties at contact zones. There were no differences in genetic diversity or population structure between dimorphic and trimorphic populations, and chloroplast haplotypes were not associated with either sexual system, indicating transitions in morph structure within each maternal lineage. M-morph frequencies were positively correlated with differentiation at microsatellite loci, indicating that the evolutionary processes influencing these neutral markers also influence alleles controlling the style morphs.     

Assessment of the domestication state of ackee (Blighia sapida K.D. Koenig) in Benin based on AFLP and microsatellite markers

Ackee (Blighia sapida) is a native multipurpose species important for the livelihoods of the rural populations in Benin. Trees are found in natural forests or are managed by farmers in different traditional agroforestry systems. Genetic variation at amplified fragment length polymorphism (AFLP) markers, four nuclear microsatellites (nSSRs) and one chloroplast microsatellite (cpSSR) were investigated in 279 individuals from six wild and eight cultivated populations from Benin. The AFLP data revealed moderate levels of diversity of ackee in Benin (mean diversity values are proportion of polymorphic loci = 52.8% and Nei??s gene diversity = 0.157, for 375 AFLP fragments). The mean diversity values based on nSSR-markers are expected heterozygosity = 0.286, allelic richness = 2.77. Genetic variation of wild and cultivated populations did not differ markedly. AMOVA revealed that only 7.3 and 5.2% of the variation was partitioned among populations for nSSR- and AFLP-markers, respectively. A Mantel test based on these both marker-types revealed significant correlations between population pairwise geographic distance and genetic differentiation. Differentiation among cultivated populations was higher than among wild populations. The only polymorphic chloroplast microsatellite marker (ccmp7) showed three haplotypes. Cultivated populations from northeastern Benin were fixed on one haplotype which was not observed elsewhere indicating a different origin of these populations possibly from neighboring Nigeria. Farmer-led domestication had an impact on the spatial distribution of genetic variation but did not result in significant losses of diversity within populations. Measures to conserve genetic resources of ackee in each of the three main bioclimatic zones in Benin are proposed.     

Phenotypic and allozyme variation in Mediterranean and desert populations of wild barley,Hordeum spontaneum Koch

Populations of wild barley, Hordeum spontaneum Koch, were collected in two distinct climatic regions, desert and Mediterranean. Plants from five desert and five Mediterranean populations were compared and contrasted for extent and structure of phenotypic variation. These same 10 and one other population from each region were analyzed for allozyme variation. In a field trial of phenotypic diversity, two phenological and 14 morphological traits were examined. Study of allozyme variation was performed using eight enzyme systems encoding for 13 loci. Plants from the desert and Mediterranean regions were significantly different in seven of 16 phenotypic traits, exhibited a high (30%) interregional component of phenotypic variation, and showed a high degree of segregation on a principal component scattergram indicating ecotypic differentiation. Mediterranean populations were twice as variable as desert populations in reproductive growth parameters (stem and spike length) and grain filling (spikelet weight), but half as variable for onset of reproduction. The extent and structure of phenotypic and allozyme variation did not match. The Mediterranean and desert populations did not differ in amount of allozyme variation as estimated by mean number of alleles per locus, effective number of alleles, polymorphism, and gene diversity (n(a), n(e), P, and H(e)), did not segregate on the basis of population genetic distances, and exhibited a low proportion of interregion allozyme diversity (2%). No effect of selection on allozyme distribution was detected. Our results suggest that the adaptation of plants originating from desert and Mediterranean environments is reflected in phenotypic but not in allozyme variation.     

Impact of human management on the genetic variation of wild pepper, Capsicum annuum var. glabriusculum

Management of wild peppers in Mexico has occurred for a long time without clear phenotypic signs of domestication. However, pre-domestication management could have implications for the population's genetic richness. To test this hypothesis we analysed 27 wild (W), let standing (LS) and cultivated (C) populations, plus 7 samples from local markets (LM), with nine polymorphic microsatellite markers. Two hundred and fifty two alleles were identified, averaging 28 per locus. Allele number was higher in W, and 15 and 40% less in LS and C populations, respectively. Genetic variation had a significant population structure. In W populations, structure was associated with ecological and geographic areas according to isolation by distance. When LM and C populations where included in the analysis, differentiation was no longer apparent. Most LM were related to distant populations from Sierra Madre Oriental, which represents their probable origin. Historical demography shows a recent decline in all W populations. Thus, pre-domestication human management is associated with a significant reduction of genetic diversity and with a loss of differentiation suggesting movement among regions by man. Measures to conserve wild and managed populations should be implemented to maintain the source and the architecture of genetic variation in this important crop relative.     

Toxic hydrogen sulfide and dark caves: phenotypic and genetic divergence across two abiotic environmental gradients in Poecilia mexicana

Divergent natural selection drives evolutionary diversification. It creates phenotypic diversity by favoring developmental plasticity within populations or genetic differentiation and local adaptation among populations. We investigated phenotypic and genetic divergence in the livebearing fish Poecilia mexicana along two abiotic environmental gradients. These fish typically inhabit nonsulfidic surface rivers, but also colonized sulfidic and cave habitats. We assessed phenotypic variation among a factorial combination of habitat types using geometric and traditional morphometrics, and genetic divergence using quantitative and molecular genetic analyses. Fish in caves (sulfidic or not) exhibited reduced eyes and slender bodies. Fish from sulfidic habitats (surface or cave) exhibited larger heads and longer gill filaments. Common-garden rearing suggested that these morphological differences are partly heritable. Population genetic analyses using microsatellites as well as cytochrome b gene sequences indicate high population differentiation over small spatial scale and very low rates of gene flow, especially among different habitat types. This suggests that divergent environmental conditions constitute barriers to gene flow. Strong molecular divergence over short distances as well as phenotypic and quantitative genetic divergence across habitats in directions classic to fish ecomorphology suggest that divergent selection is structuring phenotypic variation in this system.     

The Structure of Morphological and Genetic Diversity in Natural Populations of Carica papaya (Caricaceae) in Costa Rica

In Costa Rica, dioecious Carica papaya has been observed growing in disturbed areas and within secondary lowland forests. Such populations can serve as a reservoir of genetic and morphological diversity for this tropical fruit crop. We quantify the levels and patterns of the diversity of naturally occurring populations of C. papaya and address the demographic history of these populations. We measured 29 vegetative and reproductive morphological traits in situ from 252 plants and found significant heterogeneity among regional populations in the majority of these traits. Significant variation was found among regional populations with respect to fruit size and shape, with plants in the Nicoya Peninsula possessing smaller, less fleshy fruit, a characteristic of previously described wild populations of papaya. We then assessed the levels and patterns of genetic diversity in 164 plants from natural populations and 20 cultivars. Natural populations exhibit a deficiency of heterozygotes; however, this is much more pronounced within the cultivars. Although there is little genetic differentiation among natural populations, we did find evidence of cryptic genetic population structure. Analyses of population demography indicate that these natural populations have undergone a recent genetic bottleneck, followed by recent population expansion, possibly promoted by the transformation of the Costa Rican landscape for agricultural use. Abstract in Spanish is available at http://www.blackwell‐synergy.com/loi/btp .     

Heterogeneous selection on trichome production in Alaskan Arabidopsis kamchatica (Brassicaceae)

? Premise of the study: Environmental heterogeneity is thought to be one of the primary factors in the evolutionary maintenance of morphological variation. Here, we explore the role of environmental heterogeneity in the maintenance of variation in leaf hair (trichome) production in Arabidopsis kamchatica. ? Methods: We investigate abiotic correlates of trichome production in A. kamchatica via surveys of both herbarium specimens and wild populations. In addition, we examine patterns of phenotypic selection on trichome production among populations that differ in environmental characteristics. ? Key results: Trichome-producing herbarium specimens were more likely to occur at lower latitudes and in locations with lower mean annual precipitation and less annual variation in temperature than glabrous specimens. In surveys of wild populations, frequencies of trichome-producing plants were higher in drier habitats than in wetter environments. Using phenotypic selection analysis, we found divergent selection through female fitness (fruit production) on trichome number in populations that differ in environmental characteristics; there was selection for reduced trichome number in one population and selection for increased trichome number in another population. In a population containing both glabrous and trichome-producing plants, glabrous plants produced significantly more fruits than trichome-producing individuals, which indicates selection against the trichome morph. ? Conclusions: Our results demonstrate that there is heterogeneity in selection among populations, which could be responsible for the maintenance of trichome variation in Alaskan populations of A. kamchatica.