VARIATION IN POLLEN FLOW WITHIN AND AMONG POPULATIONS OF IPOMOPSIS AGGREGATA

Pollen dispersal is a major component of gene flow in plant populations. It can influence microevolution within and among populations as well as the evolution of floral characters that affect dispersal. Most previous studies have relied on point estimates to characterize dispersal distances, even though there is likely to be substantial intrapopulational and interpopulational variation. We measured variation in pollen dispersal for the hummingbird-pollinated herb Ipomopsis aggregata (Polemoniaceae), using powdered fluorescent dyes to estimate pollen movement. Analysis of 5–6 natural populations in each of three years indicated that mean and mean squared distances of pollen dispersal, measured over the reproductive lifespan of individual plants, varied more than threefold among populations and years. Dispersal distances also shifted over the season within a given population. Unlike the variation among populations, these seasonal changes were associated in part with changes in flower density. The mean distance of pollen dispersal from an individual plant was unrelated to the date of first flowering, but did reflect two floral characters. Plants with higher variance in stamen length across flowers delivered pollen farther on average, as predicted by computer simulations of pollen carryover. Plants with lower mean stamen lengths also delivered pollen farther. Such effects of plant characters on pollen dispersal are a critical prerequisite for dispersal to evolve in response to its effects on fitness.     

Symbiotic status alters fungal eco-evolutionary offspring trajectories

Despite host-fungal symbiotic interactions being ubiquitous in all ecosystems, understanding how symbiosis has shaped the ecology and evolution of fungal spores that are involved in dispersal and colonization of their hosts has been ignored in life-history studies. We assembled a spore morphology database covering over 26,000 species of free-living to symbiotic fungi of plants, insects and humans and found more than eight orders of variation in spore size. Evolutionary transitions in symbiotic status correlated with shifts in spore size, but the strength of this effect varied widely among phyla. Symbiotic status explained more variation than climatic variables in the current distribution of spore sizes of plant-associated fungi at a global scale while the dispersal potential of their spores is more restricted compared to free-living fungi. Our work advances life-history theory by highlighting how the interaction between symbiosis and offspring morphology shapes the reproductive and dispersal strategies among living forms.     

Butterfly species differing in mobility show different structures of dispersal‐related syndromes in the same fragmented landscape

Mobility varies strongly between and within species, reflecting different dispersal strategies. Within species, such differences can imply suites of traits associated in syndromes. Different syndrome structures have been found within species among populations differing in the selective pressures they are exposed to. Similarly, we expect species differing in mobility to show different syndrome structures in response to similar selective pressures such as landscape fragmentation. Using butterflies originating from the same fragmented landscape, we investigated the differences in mobility syndrome between four common butterflies (Pyronia tithonus, Pararge aegeria Maniola jurtina, Pieris rapae) known to differ in their mobility. We expected individuals from the less mobile species to display a resident strategy because of high dispersal cost in this fragmented landscape, and individuals from the more mobile species to display a larger range of movement strategies. Moreover, as syndromes can only be detected whenever individuals differ in their dispersal strategies, we expected mobility syndromes to be observable only in populations where dispersal polymorphism is maintained. We thus expected stronger correlations between mobility‐related traits in more mobile species. Using three mobility tests in controlled conditions designed to measure different components of mobility, we showed that mobility‐related traits were indeed correlated only in the most mobile species. The absence of correlation in the less mobile species may be explained by a low variation in movement strategies, dispersal being counter‐selected.     

ORIGINAL ARTICLE: Cycads in the insular South‐west Pacific: dispersal or vicariance?

Aim Cycads constitute an ancient plant group that is generally believed to disperse poorly. However, one group of cycads (subsection Rumphiae) is thought to have dispersed relatively recently from a Malesian source area westwards to East Africa and eastwards into the Pacific, using a floatation‐facilitating layer in their seeds. We use morphological and allozyme characters to investigate the relationships among the species within this group and to deduce whether the wide distribution was achieved by recent dispersal (as evidenced by high genetic similarity) or more distant vicariance events (high genetic differentiation). Location We examined specimens collected throughout the range of subsection Rumphiae, from East Africa through Southeast Asia to Tonga in the South‐west Pacific. Methods We investigated relationships within subsection Rumphiae of the genus Cycas by analysing 18 variable (11 informative) morphological characters and 22 allozyme loci for seven of the 10 species currently assigned to this taxon. Results Distinctive morphological characters are few and fail to resolve relationships within the group. Allozyme data show that species within this subsection are closely related and suggest that there are two groups within the subsection, one comprising Cycas thouarsii (East Africa) and C. edentata (the Philippines), and the other the remaining species (from Malesia and the Pacific). The Australian species C. silvestris is sister to subsection Rumphiae in the morphological analysis but is closely allied to C. rumphii (nested within the subsection) in the allozyme analysis, suggesting that Rumphiae may be paraphyletic and that characters thought to be taxonomically important may need to be re‐evaluated. Main conclusions Cycads within subsection Rumphiae are closely related, and the wide distribution of this group was probably achieved through relatively recent oceanic dispersal events. Separate events probably account for the dispersal of these cycads into the Pacific and to Africa. The origin and distribution of C. silvestris (Australia) could be explained by a dispersal event from New Guinea or may have resulted from a former land connection between Australia and New Guinea.     

Temporal and spatial variations in microclimate influence the larval foraging behaviors and performance of a conifer-feeding sawfly

Herbivorous insects are often exposed to broad temporal and spatial variations in microclimate conditions within their host plants and have adapted a variety of behaviors, such as avoidance or basking, to either offset or benefit from such variation. Field experiments were carried out to investigate the influence of daily and intratree variations in microclimate on the behaviors (feeding, resting, dispersal, and hiding) and associated performance of late-instar larvae of the yellowheaded spruce sawfly, Pikonema alaskensis (Rohwer) (Hymenoptera: Tenthredinidae) within crowns of 1.25-1.5 m tall black spruce (Picea mariana [Miller] Britton Sterns Poggenburg); late instars feed on developing shoots of young spruce and are often exposed to microclimatic extremes with unknown effects on performance. Larvae fed diurnally from just after dawn (0800 h) until dusk (2000 h) and rested throughout the night, with brief periods of dispersal occurring in the morning and evening. Neither larval behavior nor abiotic conditions differed significantly between the upper and lower crowns of trees. Temperature, humidity, and solar insolation all explained >90% of variation in feeding; however, sunrise and sunset were the most likely cues influencing diurnal behavior. Most larvae (94%) fed on the bottom, shaded side of shoots, and field experiments indicated that this behavior is adaptive with respect to microclimate, probably reducing hygrothermal stress. Thus, behavioral adaptations by P. alaskensis to daily and within-shoot microclimatic variation may reduce the risk of hygrothermal stress during dispersal or feeding, while still allowing larvae to feed on the preferred and highly nutritious upper crown foliage of young spruce.     

动物对孢子植物的传播模式及进化意义

孢子植物物种多样性丰富, 是自然生态系统的重要组成部分。孢子植物的传播通常被认为主要依靠风、水、弹力等非生物媒介, 而动物的作用往往被忽略。本文主要概述了: (1)孢子植物对动物传播的适应: 一方面孢子植物可为动物提供食物、庇护所、繁殖场所等, 另一方面孢子植物也可产生视觉、嗅觉等方面的线索来吸引动物, 从而促进动物传播其繁殖体。(2)动物对孢子植物的传播模式: 包括体内传播(消化道和组织寄生)和体外传播两种, 这些模式都能对孢子植物繁殖体进行有效传播。由于动物间形态或生活习性的不同, 以致传播距离存在差异, 最短距离为0.1 cm, 最长距离可从北半球至南半球。(3)动物对孢子植物传播的生态与进化意义; 由于某些孢子植物繁殖体的结构特点或萌发的需求, 以致其繁殖体只能通过动物的传播才能得以定殖, 因此动物与孢子植物之间存在密不可分的关系。目前, 动物对孢子植物的传播研究主要是描述性的内容以及研究单方面的传播途径, 建议在今后的研究中考虑动物对孢子植物传播的有效性以及多途径同时传播对孢子植物定殖的影响, 同时应更加关注孢子植物和动物互惠关系的形成、维持机制及将来的进化趋势。     

Spore deposition of wood-decaying fungi: importance of landscape composition

The abundance of many species of wood-decaying fungi has decreased dramatically in Swedish boreal forests over the last century. Therefore, we have investigated the relationship between the spore dispersal of wood-decaying fungi and two key features of landscape composition, namely the amount and age of old forest stands at different spatial scales. Spore deposition was monitored in two regions, using sampling methods based on recording the dikaryotisation of monokaryotic mycelia on nutrient agar and wood discs. The studied species, all mainly confined to Norway spruce in the boreal forest zone, were Fomitopsis pinicola, Fomitopsis rosea , Gloeoporus taxicola , Phlebia centrifuga and Trichaptum laricinum . The study of forestry intensity showed for both regions, that the spore deposition for F. pinicola , F. rosea and G. taxicola was higher in circular plots (radius 2-km) with a high proportion of old Norway spruce forest (>80 yr) than in plots with a lower proportion of old forest. Analysis of the variation in spore deposition of F. rosea and P. centrifuga in relation to the proportion of old spruce forest within 1, 2 and 3 km of the spore sampling point showed that the proportion of old forest within a 3-km radius explained more of the variation than the proportions within 1- and 2-km radii. In addition, the proportion of forest older than 140 yr explained more of the variation than the proportion of younger forests. Thus, the results show that the spore deposition of the studied species strongly depends on the landscape composition at both regional and local scales. Further, the spore deposition at the local scale was best explained by the proportion of >140 yr old spruce forest, which exceeds the common harvest rotation period.     

Correlates of diversification in the plant clade Dipsacales: geographic movement and evolutionary innovations

We explore patterns of diversification in the plant clades Adoxaceae and Valerianaceae (within Dipsacales), evaluating correlations between biogeographic change (i.e., movements into new areas), morphological change (e.g., the origin of putative key innovations associated with vegetative and reproductive characters), and shifts in rates of diversification. Our findings indicate that rates of diversification in these plants tend to be less tightly correlated with the evolution of morphological innovations but instead exhibit a pronounced correlation with movement into new geographic areas, particularly the dispersal of lineages into new mountainous regions. The interdependence among apparent novelties (arising from their nested phylogenetic distribution) and the correlation between morphological and biogeographic change suggests a complex history of diversification in Dipsacales. Overall, these findings highlight the importance of incorporating biogeographic history in studies of diversification rates and in the study of geographic gradients in species richness. Furthermore, these results argue against a simple deterministic relationship between dispersal and diversification: like other factors that may influence the probability of speciation and/or extinction, the impact of dispersal on diversification rates depends on being in the right place at the right time.     

大小蠹来源地的COⅠ基因标记

【背景】大小蠹属昆虫是重要的林木害虫,我国口岸有多次截获记录,确定大小蠹的来源地可以有针对性地加强对大小蠹的检验检疫工作。【方法】测定了5种高风险大小蠹(红脂大小蠹、红翅大小蠹、中欧山松大小蠹、落叶松大小蠹和间大小蠹)共12个样本的线粒体DNA细胞色素氧化酶C亚基Ⅰ基因(COⅠ)的部分序列。【结果】利用巢式PCR技术,在不同种类的大小蠹样本中均获得了530bp的靶标片段,比对分析显示,不同种大小蠹之间的COⅠ序列差异显著,同种大小蠹不同个体之间的COⅠ序列存在一定的差异,但差异不显著。系统进化树分析结果显示,大小蠹可以明显分为2支,其中,间大小蠹单独为一分支,另一分支由红脂大小蠹、红翅大小蠹、中欧山松大小蠹和落叶松大小蠹组成;同时,同种大小蠹不同个体的来源情况在进化树中有一定体现。【结论与意义】COⅠ基因可以较好地反映样本的来源地,对今后制定大小蠹的具体检验检疫措施有实际指导意义。     

The adaptive background of nannandry: dwarf male distribution and fertilization in the moss Homalothecium lutescens

Dwarf males (nannandry) occur in many unrelated, mostly aquatic, groups of organisms. Among land plants they are unique to bryophytes. In this study our aim was to explain variation in frequency of dwarf males and fertilization within populations in the moss Homalothecium lutescens. We compared parameters related to dwarf male presence and sporophyte production in 90 colonies from three localities. Dwarf male density was positively associated with colony moisture at two of the localities, suggesting increased spore germination and dwarf male survival with moist conditions. At one of these localities, dwarf male density was also positively associated with the presence of perichaetia (female sexual branches). Dwarf male density and fertilization frequency were positively associated in two of the localities. Furthermore, in one population, fertilization was also positively associated with canopy cover, which could be attributed to improved nutrient status as a result of throughfall, increased sperm‐dispersal efficiency because of larger water drops, or more favourable moisture conditions as a result of shading. Nannandry thus appears to strongly reduce the problem of short fertilization distances in bryophytes, but the presence of water is still critical because the dwarf males are dependent on a certain level of humidity for recruitment and/or development. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113 , 74–84.     

Population genetics of the carinate pillsnail, <Emphasis Type="Italic">Euchemotrema hubrichti</Emphasis>: genetic structure on a small spatial scale

Euchemotrema hubrichti (Pilsbry HA (1940) Volume 1, Part 2. Monographs of the Academy of Natural Sciences, Philadelphia, 3, I-VI, 2, 575–994, I-IX) is a polygyrid land snail known only from the Larue Pine Hills region of southwestern Illinois, USA. Despite the restricted range of the species, observations in the field suggested that patterns of genetic variation within E. hubrichti might show geographic structure. To assess the dispersal potential of E. hubrichti, a mark-recapture experiment was performed at two sites <100 meters apart. To evaluate the geographic distribution of genetic variation, a region of the mitochondrial cytochrome c oxidase subunit I gene was sequenced from 53 snails collected from sites spanning the species range. Mark-recapture experiments suggested that population sizes at two sites were low and migration between sites was not detected. AMOVA and nested clade analysis (NCA) revealed significant genetic structure among sites. Two sites in particular appeared to be isolated from the others based on nested clade analysis. AMOVA also detected significant genetic differentiation among sites, but partitioning the data into two groups (populations 1 and 2) based on the NCA results did not explain a significant amount of genetic variation. Coalescent methods suggested that population sizes and migration rates for populations 1 and 2 are unequal. These patterns are likely due to the low dispersal ability of E. hubrichti and patchiness of preferred E. hubrichti habitat rather than an ancient vicariant event. These preliminary findings have implications both for the conservation of this species and for the population genetics of any spatially restricted endemic species with low dispersal ability.     

Consequences of pre-dispersal damage by insects for the dispersal and recruitment of mangroves

Herbivores may enhance plant recruitment, but such positive interactions may be overlooked in favour of obvious negative effects of herbivory on propagules. My objective was to determine whether larval insects that feed and develop within fruit of the mangrove Avicennia marina act as mutualist herbivores by increasing the dispersal of propagules without affecting their viability and emerging successfully as adults following dispersal of the propagule by water. Surveys revealed that frugivory is common throughout the mangrove forest, and fruit had up to six exit holes where larvae had emerged as adults. Larval insects did not affect the flotation of propagules with pericarps, a thin structure that provides buoyancy for dispersal by water. In contrast, after simulating germination by removing the pericarp, the majority of propagules with three exit holes floated on average for 20 h longer than those without exit holes, which sank immediately. Based on this evidence that frugivory could increase the dispersal potential of propagules, I predicted that propagules consumed by larval insects should disperse farther than undamaged propagules, and this was tested by quantifying the potential viability of propagules stranded on beaches at increasing distances (up to 20 km) from mangrove forests. Flies and moths emerged as adults after being transported tens of kilometres within mangrove propagules, revealing a novel mode of dispersal. Proportionally fewer potentially viable propagules were supplied to beaches at increasing distances from mangrove forests, however, indicating that larval insects negatively affect recruitment and are thus not acting as mutualist herbivores. Nevertheless, when transported back to the mangrove forest, seedlings established from propagules damaged by larval insects and stranded on beaches. Therefore, although frugivory does not preclude mangrove recruitment, its negative effects in the pre-dispersal environment may be intensified with increasing dispersal distance, thus limiting the long-distance supply of propagules and recruitment of mangroves.     

Does fruiting phenology vary with fruit syndrome? An investigation on animal-dispersed tree species in an evergreen forest in south-western Cameroon

Fruit syndrome found in zoochorous plants is regarded as a result of hypothetical coevolution with a seed disperser/predator. Fruiting phenology was compared among two representative syndromes, such as bird–monkey syndrome (BM) and ruminant–rodent–elephant syndrome (RRE), plus the gravity dispersal species for comparison, in south-western Cameroon in order to examine which biotic or abiotic factors educed syndrome variation. The individual size of selected plants (> 1.6 m in height) was recorded in a 16.95-ha area for recruitment estimation and their fruiting behaviour was checked for 9 months from June 1985 to February 1986. The BM species, a suggested successful group, fruited in a less synchronous manner within species and had fruiting peaks just before and during the rainy season. The RRE species, a suggested failing group, showed greater variation within syndrome in fruiting timing, duration and synchronization. Results obtained supported no clear phenological response to corresponding vertebrates except for the case of Sacoglottis gabonensis in the RRE. The reason for concealing potential responses is probably a result of conflicting requirements for seed dispersal and offspring survival. Periodical fruiting of the BM is likely to reflect ecological constraint, such as water stress on seedlings, caused from syndrome-specific morphological limitation. In this case, the animals have indirect effects on fruiting phenology through selecting syndrome-specific fruit morphology. The greater variation of the RRE suggests a broad spectrum of dispersal tactics from faithful zoochory to alternatives, with a trade off between agent restriction and seed size and another trade off between parental investment and seed-predation risk.     

Gynoecium, fruit and seed structure of Paullinieae (Sapindaceae)

Despite an emphasis on fruit characters in Paullinieae taxonomy, few detailed morphological and anatomical studies of the gynoecia, fruits and seeds exist. The aims of the present study were (1) to provide a detailed documentation of gynoecium, fruit and seed structure and ontogeny in selected Paullinieae taxa; (2) to determine whether the gynoecium, seed and seedling provide additional characters of systematic value within the tribe; and (3) to relate the structural findings to mechanisms of fruit dehiscence and dispersal within these taxa. Newly described characters of systematic value within Paullinieae are shape and surface of the obturator, type of pollen tube transmitting tract, indumentum of the inner and outer surface of the carpels, ovary wall anatomy, aril anatomy, pseudo-hilum form, seedling germination mode and structure of first leaves. The fruits of Paullinia are septifragal, and conspicuous colour contrasts between the pericarp, aril and seed in most species of this genus are suggestive of a bird dispersal syndrome. Interestingly, it appears that relatively minor structural changes are associated with switches to rodent dispersal in Paullinia sphaerocarpa and water dispersal in P. clathrata and P. hystrix. Anemochorous fruits are septifragal ( Cardiospermum and Urvillea ) or schizocarpic ( Houssayanthus , Lophostigma , Serjania ). They are structurally similar and Cardiospermum with septifragal capsules may also show septicidal dehiscence.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 147 , 159–189.     

Dispersal of spores and pollen from crops

Fungal spores and pollens can be dispersed in a number of ways: by animals and insects; by water; by wind or by rain. This paper concentrates on the effects of wind on the dispersal of spores and pollen grains and the effects of rain on spore dispersal. For dispersal to be successful particles must complete three phases: removal, dispersal through the air and deposition. The biology of the organism and its environment can affect all three phases, however, once released the fate of all airborne particles largely depends on the laws of physics which govern the motion of the air. Many types of spore are actively ejected into the air while others are simply blown from the host surface. Particle size and shape affects dispersal and deposition phases. Local environmental factors such as temperature, humidity and light, as well as wind or rain, can play a key role in the removal of spores. Wind speed and turbulence or rainfall, largely determine spore dispersal, but, the size and shape of the particle, the nature of the plant canopy and the way the particles are released into the air may also be important. Particle deposition depends on both environmental and biological factors. This paper briefly considers these processes using examples and how they can be modelled.     

Evolutionary syndromes linking dispersal and mating system: the effect of autocorrelation in pollination conditions

Self-fertilization is classically thought to be associated with propagule dispersal because self-fertilization is a boon to colonizers entering environments devoid of pollinators or potential mates. Yet, it has been theoretically shown that random fluctuations in pollination conditions select for the opposite association of traits. In nature, however, various ecological factors may deviate from random variations, and thus create temporal correlation in pollination conditions. Here, we develop a model to assess the effects of pollination condition autocorrelation on the joint evolution of dispersal and self-fertilization. Basically, two syndromes are found: dispersing outcrossers and nondispersing (partial) selfers. Importantly, (1) selfers are never associated with dispersal, whereas complete outcrossers are, and (2) the disperser/outcrosser syndrome is favored (resp. disfavored) by negative (resp. positive) autocorrelation in pollination conditions. Our results suggest that observed dispersal/mating system syndromes may depend heavily on the regime of pollination condition fluctuations. We also point out potential negative evolutionary effects of anthropic management of the environment on outcrossing species.     

Genetic variation of morphological characters within a single isolate of the endomycorrhizal fungus Glomus clarum (Glomaceae)

The nature of variation in morphological characters in spores of arbuscular endomycorrhizal fungi (Order Glomales, Class Zygomycetes) has received little attention, despite the importance of these characters in modern taxonomy of the order. We tested the hypothesis that genetic variation exists in spore size and color (presumably important taxonomic characters) within a single isolate of the glomalean fungus Glomus clarum. Phenotypic variation in size and color of spores was determined from a pot culture population (designated P). A 10% selection pressure was imposed on replicate pot cultures of the first progeny culture generation (G1) by selecting the smallest, largest, yellowest, and whitest spores from the P generation and inoculating Sorghum bicolor plants. The experiment was repeated for another generation (G2), but with a 5% selection pressure. In both the G1 and G2 generations, significant differences in spore size and color were observed among the various treatments, indicating substantial genetic variation in these characters. Despite efforts to keep the physical environment constant across generations, we observed variation in the overall means of spore size and color among the generations (regardless of treatment), indicating a strong nongenetic influence on character expression. This study provides empirical evidence that will help delimit species boundaries among isolates of Glomus clarum and similar morphospecies. It also demonstrates a promising method to help elucidate the nature of character diversity in obligately asexual fungi.     

Fruit characters as a basis of fruit choice and seed dispersal in a tropical forest vertebrate community

Summary Interactions between a large community of vertebrate frugivore-granivores (including 7 species of large canopy birds, 19 species of rodents, 7 species of ruminants, and 6 species of monkeys), and 122 fruit species they consume, were studied for a year in a tropical rainforest in Gabon.The results show how morphological characters of fruits are involved in the choice and partitioning of the available fruit spectrum among consumer taxa. Despite an outstanding lack of specificity between fruit and consumer species, consideration of simple morphological traits of fruits reveals broad character syndromes associated with different consumer taxa. Competition between distantly related taxa that feed at the same height is far more important than has been previously supposed. The results also suggest how fruit characters could have evolved under consumer pressure as a result of consumer roles as dispersers or seed predators. Our analyses of dispersal syndromes show that fruit species partitioning occurs more between mammal taxa than between mammals and birds. There is thus a bird-monkey syndrome and a ruminant-rodent-elephant syndrome. The bird-monkey syndrome includes fruit species on which there is no pre-dispersal seed predation. These fruits (berries and drupes) are brightly colored, have a succulent pulp or arillate seeds, and no protective seed cover. The ruminant-rodent-elephant syndrome includes species for which there is pre-dispersal predation. These fruits (all drupes) are large, dull-colored, and have a dry fibrous flesh and well-protected seeds.     

Phylogeographic structure and cryptic speciation in the trans-Antarctic moss Pyrrhobryum mnioides

Abstract Many bryophyte species have distributions that span multiple continents. The hypotheses historically advanced to explain such distributions rely on either long-distance spore dispersal or slow rates of morphological evolution following ancient continental vicariance events. We use phylogenetic analyses of DNA sequence variation at three chloroplast loci ( atpB-rbcL spacer, rps4 gene, and trnL intron and 3'spacer) to examine these two hypotheses in the trans-Antarctic moss Pyrrhobryum mnioides. We find: (1) reciprocal monophyly of Australasian and South American populations, indicating a lack of intercontinental dispersal; (2) shared haplotypes between Australia and New Zealand, suggesting recent or ongoing migration across the Tasman Sea; and (3) reciprocal monophyly among Patagonian and neotropical populations, suggesting no recent migration along the Andes. These results corroborate experimental work suggesting that spore features may be critical determinants of species range. We use the mid-Miocene development of the Atacama Desert, 14 million years ago, to calibrate a molecular clock for the tree. The age of the trans-Antarctic disjunction is estimated to be 80 million years ago, consistent with Gondwanan vicariance, making it among the most ancient documented cases of cryptic speciation. These data are in accord with niche conservatism, but whether the morphological stasis is a product of stabilizing selection or phylogenetic constraint is unknown.