A multifaceted ecological assessment reveals the invasion of the freshwater red macroalga Montagnia macrospora (Batrachospermales,Rhodophyta) in Taiwan

Invasive freshwater macroalgae are rarely described. Montagnia macrospora is a freshwater red alga introduced from South America to East Asia via the global aquarium trade. The earliest occurrence record of this alga in Taiwan is dated 2005. To determine whether M. macrospora has become invasive in Taiwan and to understand the traits that facilitated its invasion, we took a multifaceted approach that combines examination of ecological background and population genetic analysis. Our island‐wide survey showed that M. macrospora is widespread in the field across Taiwan, where the climate greatly differs from that of South America, and can self‐sustain for nearly a decade. Our population genetic analysis revealed a lack of genetic diversity of M. macrospora in Taiwan, consistent with the hypothesis that the alga expanded through asexual reproduction. Moreover, during our long‐term ecological assessments and field surveys, we observed that M. macrospora is an ecological generalist that can survive in a wide range of temperature, pH, illumination, and nutrient enrichment. Taken together, our data suggest that M. macrospora has successfully invaded the freshwater ecosystems of Taiwan, likely due to its ability to disperse asexually and to grow under broad environmental conditions. We hope that our study brings attention to invasive freshwater algae, which have been overlooked in conservation planning and management.     

东亚与北美当归属花粉形态的比较研究

对东亚与北美当归属（ＡｎｇｅｌｉｃａＬ．）７２种植物的花粉形态特征进行了观察,表明当归属的花粉为一多类型的类群,包含伞形科全部６种花粉类型,说明该属既是一个古老的属,同时又是在发展中不断演化的属。东亚和北美的共有种表明两地区的当归属十分近缘。从两地区存在着古老的姊妹群推断它们共同起源于白垩纪到老第三纪。中国的四川和北美的西北部同为当归属的起源和演化中心。花粉特征还表明当归属的近缘属山芹属（ＯｓｔｅｒｉｃｕｍＭａｘｉｍ．）为一自然的分类群     

Reduction in the number of Early Cambrian helicoplacoid species

Helicoplacoid echinoderms occur in Lower Cambrian shales and mudstones along the northern coast of Laurentia, now western North America. The group originally contained four genera and nine species, diagnosed by differing ambulacra, plate length-to-width ratios, and plate ornament. Ambulacra are identical in all specimens except for the single example of Waucobella nelsoni, and the remaining morphologic differences are caused by ontogeny. The characters used to diagnose seven of the original nine species are therefore invalid, reducing the group to three genera and three species. These are: Helicoplacus gilberti Durham and Caster, 1963, Waucobella nelsoni Durham, 1967, and Polyplacus kilmeri Durham, 1967, which has a test composed of ambulacra, and likely represents a developmental oddity.     

Latest Helcionelloid Molluscs from the Lower Ordovician of Kazakhstan

The helcionelloid mollusc Chuiliella elenae gen. et sp. nov. is described from the Lower Ordovician of Kazakhstan. It represents the geologically youngest record of a group of mainly bilaterally symmetrical ancient molluscs which originated in the earliest Cambrian, flourished during the early–mid Cambrian and was thought to have become extinct during the late Cambrian. Chuiliella is a typical helcionelloid in terms of shell shape, although the comarginal ornamentation characteristic of many helcionelloids is lacking. Interpretation of the raised margin of the aperture adjacent to the earlier coiled whorl as exhalant channels favours reconstruction of helcionelloids as endogastrically coiled, i.e., with the apex posterior.     

Problematic cap-shaped fossils from the Lower Cambrian of North-East Greenland

Three species ofOcruranus Liu, 1979 are described from the Bastion Formation of North-East Greenland, of late Early Cambrian (middle Dyeran of North American usage) age, representing the youngest record of a genus originally described from the earliest Cambrian Meishucunian Stage of China. An accompanying species, tentatively assigned toXianfengella He &Yang, 1982, seems also to be present in South Australia in strata of late Early Cambrian (Botoman of Siberian usage) age, although also this genus was described originally from the Meishucunian.Ocruranus andXianfengella from China have been interpreted as possibly parts of coeloscleritophoran scleritomes, perhaps halkieriids, rather than individual molluscan shells. Their shell form is not typical of helcionelloids which otherwise dominate the Early Cambrian molluscan record, but conclusive evidence of affinity is not forthcoming from the Greenland records. New taxa:Ocruranus septentrionalis n. sp. andOcruranus tunuensis n. sp.      

A new leptarctine (Carnivora: Mustelidae) from the early Miocene of the northern Tibetan Plateau: implications for the phylogeny and zoogeography of basal mustelids

A recently discovered basal mustelid carnivoran, Kinometaxia guangpui gen. et sp. nov. , is described from the lower Miocene Tiejianggou Formation in the Danghe (Tabenbuluk) area of the northern Tibetan Plateau, north-western Gansu Province, China. The new Chinese form has a double temporal crest and roughened surface on the temporal area, a small orbit, a deep zygomatic arch, an anterolaterally expanded ectotympanic that fuses with the postglenoid process, and a lateral opening of the postglenoid foramen. These cranial morphologies suggest membership in the leptarctines, a group of markedly hypocarnivorous taxa mostly confined to the Miocene of North America. The rather hypercarnivorous dentition of the new Chinese form, however, is in sharp contrast to that of the traditionally defined leptarctines that have complex and quadrate M1s and P4s. Phylogenetic analysis suggests a sister relationship of the new Chinese mustelid and Schultzogale , and together they form a basal sister clade to the advanced leptarctines such as Craterogale , Leptarctus , and Hypsoparia . The Chinese form further expands the concept of the leptarctine clade and adds additional complexity to the early evolution of the Mustelidae. It bridges a morphological gap between European and North American taxa, and supplies an important piece to the puzzle of the zoogeography of basal mustelids. Furthermore, the new leptarctine helps to constrain the age of the Danghe strata, which has important implications for the timing of tectonic activity in the northern Tibetan Plateau.  © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society , 2004, 142 , 405−421.     

Distribution and systematics of the freshwater red algal family Thoreaceae in North America

North American populations and type specimens of Thorea and Nemalionopsis were compared using multivariate morphometrics and image analysis. We continue to recognise the separation of the two genera based on spore-bearing branch-to-assimilatory filament length ratio and assimilatory filament density (≤0·3 and loose for Thorea and ≥0·6 and compressed for Nemalionopsis, respectively). Four species of Thorea were distinguished worldwide: T. hispida (syn. T. andina, T. lehmannii, T. ramosissima), T. violacea (syn. T. bachmannii, T. brodensis, T. gaudichaudii, T. okadae, T. prowsei and T. riekei), T. clavata and T. zollingeri. The former two species have variable branched, non-clavate assimilatory filaments, whereas the latter two have unbranched, clavate assimilatory filaments. Thorea hispida has copious secondary branches while T. violacea is sparsely branched. Thorea clavata is differentiated from T. zollingeri by having fewer monosporangia per cluster. Two of these species were found in North America: T. hispida in Mexico, Grenada and Jamaica, and T. violacea in Texas and Mexico. Two species of Nemalionopsis were differentiated: N. shawii with long assimilatory filaments composed of barrel-shaped cells and n. tortuosa (syn. N. shawii f. caroliniana) with short assimilatory filaments having cylindrical cells. Only N. tortuosa has been collected on the continent, from Florida, Louisiana and North Carolina.     

Redescription of †Paralycoptera wui Chang & Chou, 1977 (Teleostei: Osteoglossoidei) from the Early Cretaceous of eastern China

?Paralycoptera wui Chang & Chou, 1977 from the Early Cretaceous of China is redescribed herein through a re‐examination of the original materials, as well as observations on some newly collected specimens. The use of the peeling method has revealed much of the new or revised information on its osteology, e.g. aspects of the nasal, infraorbitals, retroarticular, preopercle, extrascapular, basihyal toothplate, epineural, pelvic fin, caudal skeleton and scales. The phylogenetic relationships of ?Paralycoptera and other osteoglossomorphs are re‐evaluated. The cladistic analysis largely agrees with the previous hypothesis that ?Paralycoptera is not a ?lycopterid, but rather a stem osteoglossoid. ?Paralycoptera is excluded from notopteroids and exhibits the following derived characters of the Osteoglossoidei: (1) palatal area behind and below orbit completely covered by infraorbitals; (2) jaw articulation under posterior portion of orbit; (3) opercle depth twice or more its width; (4) first pectoral fin ray much enlarged and long, extending posteriorly beyond origin of pelvic fin. ?Singida and ?Phareodus are regarded as different levels of osteoglossoids above ?Paralycoptera. ?Singida shares the following derived characters with ?Phareodus plus extant osteoglossoids: (1) jaw articulation posterior to orbit; (2) anterior process of hyomandibula in contact with entopterygoid; (3) subopercle small and anterior to opercle. ?Phareodus shares the following derived characters with extant osteoglossoids: (1) supraorbital and otic sensory canals connected; (2) one uroneural; (3) reticulate furrows present over entire scale.     

东亚帚菊木族及其相关类群的叶表皮形态比较

在光学显微镜和扫描电镜下,对东亚地区帚菊木族及其南美相关类群共12属31个种的叶表皮进行了观察.研究发现气孔器普遍存在于叶的下表皮,少数种的上表皮也有分布,气孔器主要为星型和无规则型两种类型.叶表皮细胞为多边形和不规则形,垂周壁平直弓形、浅波状或波浪状.气孔器表面观为椭圆形和近圆形,气孔外拱盖内缘近平滑至浅波状,其中 Macroclinidium 较为特殊,具有双层气孔外拱盖.叶表面通常光滑无蜡质或角质,部分种叶表面具薄片状或层状蜡质.研究表明,叶表皮特征在种间一致性较高,但在属间,特别是属群和亚族间存在一定差异.     

Mammalian Faunal Succession in the Cretaceous of the Kyzylkum Desert

Both metatherians and eutherians are known from the Early Cretaceous (Barremian, 125 mya; million years ago) of China, while eutherian-dominated mammalian faunas appeared in Asia at least by the earliest Late Cretaceous (Cenomanian, 95 mya). The approximately 99–93 my old (Cenomanian) Sheikhdzheili l.f. from western Uzbekistan is a small sample of only eutherians, including three zhelestids and a possible zalambdalestoid. The much better-known 90 my old (Turonian) Bissekty l.f. at Dzharakuduk in the central Uzbekistan includes 15 named and unnamed species, based on ongoing analyses. Of these, 12 are eutherians represented by at least the three groups—asioryctitheres, zalambdalestids, and zhelestids—plus an eutherian of uncertain position—Paranyctoides. Zalambdalestids and zhelestids have been argued to be related to the origin of the placental gliriforms (Euarchontoglires) and ferungulates (Laurasiatheria), respectively. Although there are four previously recognized metatherians, we believe three are referable to the deltatheroid Sulestes karakshi and the fourth, Sailestes quadrans, may belong to Paranyctoides. There is one multituberculate and one symmetrodont in the Bissekty l.f. While comparably aged (Turonian) localities in North America have somewhat similar non-therians, they have more metatherians and no eutherians. The next younger localities (early Campanian, ∼80 mya) in North America have both a zhelestid and Paranyctoides, suggesting dispersal of eutherians from Asia. At Dzharakuduk, the approximately 85 my old (late Turonian/Coniacian) Aitym l.f. is much less well known than the Bissekty l.f., but yields nearly identical taxa, with two non-therians, one metatherian, and six eutherians.     

Differential diversifications of South American and Eastern Asian disjunct genera Bocconia and Macleaya (Papaveraceae)

Bocconia (10 species) and Macleaya (2 species) are two disjunct genera between South America and eastern Asia (EAS) in the Papaveraceae offering an opportunity to compare its biogeographic history with that of the well‐known disjunction between EAS and eastern North American (ENA). Our phylogenetic analyses of the chloroplast matK and rbcL gene sequences of Ranunculales including two species of Macleaya and six species of Bocconia supported the monophyly of Bocconia, Macleaya, and Chelidonioideae to which Bocconia and Macleaya belong. Nucleotide sequences of matK, rbcL, and nrDNA ITS supported the sister relationship of Bocconia and Macleaya. Biogeographic analyses of Chelidonioideae using S‐DIVA (statistical dispersal vicariance analysis) and DEC (dispersal extinction cladogenesis) methods inferred Eurasia as the most likely ancestral area of Bocconia and Macleaya and suggested no extinction events in either Bocconia or Macleaya. This agrees with the “Out‐of‐Asia” pattern of the EAS‐ENA disjunction. Molecular dating of Ranunculales with fossil‐based calibrations showed that Bocconia and Macleaya diverged in the late Eocene and early Oligocene, which is much earlier than most EAS‐ENA disjunct taxa. The disjunction may have formed via long distance dispersal or boreotropical connections via the North Atlantic and Bering land bridges. Both Bocconia and Macleaya diversified in the late mid‐Miocene, but Bocconia has apparently experienced a greater diversification probably aided by the evolution of the bird dispersal syndrome in fruit and seed after migration to South America. The greater diversification of Bocconia is also evidenced by the diverse leaf morphology and growth habit in response to colonization in various local habitats in South America.     

The reticulated species of Chydorus (Cladocera,Chydoridae): two new species with suggestions of convergence

Of the two small, compact, dusky, reticulated species of Chydorus from Sri Lanka, one arbitrarily has been selected as the taxon to which Daday's name reticulatus henceforward will be attached, the other herein being described as new. A third species from eastern North America, also described herein as new, resembles the two Asian species in certain gross features but otherwise is very distinct. Particularly noteworthy among these differences are the structure of the labrum and of the male postabdomen and copulatory hook. The question is raised whether the gross morphological features that seem to unite the taxa or the features of the labrum and of the male that separate them are the more conservative. No unequivocal answer is provided. Because of their obvious close similarity, the two Asian species are established as a species group, the one from North America being divergent and therefore not included. The possibility of convergence is considered.The North American taxon in north Florida occurs in waterbodies with a pH less than 5 (down to 4.2), conductivity less than 40 µS, and usually with no more than a trace of dissolved color. They are naturally acidic, non-bog lakes. The number of species of chydorids and of total littoral Cladocera in them are considerably greater than found by Fryer in waterbodies of comparable acidity in England. The occurrence also of a diversity of macrophytes and fishes in these lakes indicates that pH per se is not the factor forcing the decreasing diversity associated with changes resulting from acid precipitation.     

Effects of taxon sampling on molecular dating for within‐genus divergence events,when deep fossils are used for calibration

A universal method of molecular dating that can be applied to all families and genera regardless of their fossil records, or lack thereof, is highly desirable. A possible method for eudicots is to use a large phylogeny calibrated using deep fossils including tricolpate pollen as a fixed (124 mya) calibration point. This method was used to calculate node ages within three species‐poor disjunct basal eudicot genera, Caulophyllum, Podophyllum and Pachysandra, and sensitivity of these ages to effects such as taxon sampling were then quantified. By deleting from one to three accessions related to each genus in 112 different combinations, a confidence range describing variation due only to taxon sampling was generated. Ranges for Caulophyllum, Podophyllum and Pachysandra were 8.4–10.6, 7.6–20.0, and 17.6–25.0 mya, respectively. However, the confidence ranges calculated using bootstrapping were much wider, at 3–19, 0–32 and 11–32 mya, respectively. Furthermore, deleting 10 adjacent taxa had a large effect in Pachysandra only, indicating that undersampling effects are significant among Buxales. Changes to sampling density in neighboring clades, or to the position of the fixed fossil calibration point had small to negligible effects. Non‐parametric rate smoothing was more sensitive to taxon sampling effects than was penalized likelihood. The wide range for Podophyllum, compared to the other two genera, was probably due to a high degree of rate heterogeneity within this genus. Confidence ranges calculated by this method could be narrowed by sampling more individuals within the genus of interest, and by sequencing multiple DNA regions from all species in the phylogeny.     

Large‐scale patterns of tree species richness and the metabolic theory of ecology

The metabolic theory of ecology (MTE) endeavours to explain ecosystem structure and function in terms of the effects of temperature and body size on metabolic rate. In a recent paper (Wang et al., 2009, Proceedings of the National Academy of Sciences USA, 106 , 13388), we tested the MTE predictions of species richness using tree distributions in eastern Asia and North America. Our results supported the linear relationship between log‐transformed species richness and the inverse of absolute temperature predicted by the MTE, but the slope strongly depends on spatial scale. The results also indicate that there are more tree species in cold climate at high latitudes in North America than in eastern Asia, but the reverse is true in warm climate at low latitudes. Qian & Ricklefs (2011, Global Ecology and Biogeography, 20 , 362–365) recently questioned our data and some of the analyses. Here we reply to them, and provide further analyses to show that their critiques are primarily based on unsuitable data and subjective conjecture.     

A test of the vicariance hypothesis of western North American freshwater biogeography

Aim The biogeography of western North American freshwater molluscs has traditionally been attributed to vicariance associated with late Tertiary rearrangement of landscape based on distributional evidence and the putatively limited dispersal ability of these organisms. We examined the phylogeography of a widely ranging western springsnail (Pyrgulopsis wongi Hershler, 1989) to test this hypothesis and evaluate the relative importance of vicariance and dispersal in structuring the distribution of this species. Location Southwestern Great Basin (California and Nevada), United States of America. Method Two mitochondrial genes (COI, NDI) were sequenced for 28 populations of P. wongi spanning its entire geographic range, which consists of 10 topographically closed drainage basins. We also sequenced eight closely related congeners, as well as the type species of the closely related eastern North American genus Floridobia Hershler & Thompson, 2002, which was used as the outgroup. Phylogenies based on the combined data set were obtained using several methods, and networks for each gene were generated as an additional means of examining relationships among haplotypes. Partitioning of haplotype variation was studied using amova , migration between populations was estimated using a coalescent‐based method (mdiv ), and divergence times were inferred using a locally calibrated molecular clock and mdiv . Results Pyrgulopsis wongi is subdivided into narrowly localized and widely distributed lineages that diverged in the Pleistocene, well after the inception of the contemporary regional landscape. While large Φ_ST values and the localized geographic distributions of most haplotypes imply absence or negligible contemporary dispersal of this spring‐dwelling snail, the pattern of phylogeographic structuring, presence of a few widespread haplotypes, and results of the mdiv analyses suggest geologically recent dispersal across drainage divides. Main conclusions Phylogeography of P. wongi conflicts with the traditional vicariance model as it is not structured by the contemporary landscape and is instead indicative of geologically recent dispersal. In the absence of evidence that dispersal of this species occurred through surface water connections during the relevant (Quaternary) time frame, we conjecture that spread may have instead been mediated by transport on waterfowl or via upland stream capture. The non‐concordance between phylogeography and landscape reported in this and other recent studies of Pyrgulopsis suggests that members of this diverse and imperiled genus should not be managed using an a priori, ‘watershed as conservation unit’ approach.     

Transgressive facies sequence of a high energy,wave-tide-storm-influenced shoreface: A case study of the east Frisian Barrier Islands (Southern North Sea)

Summary Published information and recent observation of physical and biogenic structures as well as grain size patterns along the East Frisian barrier islands (southern North Sea) have been synthesized to develop a facies model for a high energy, wave-tide-storm-influenced transgressive shoreface. As exemplified by Spiekeroog Island, three major shoreface facies (Upper, Central and Lower) are distinguishable, each of which can be subdivided into a Proximal and a Distal subfacies. A transgressive shoreface sequence in the rock record, in ascending order, will show the Upper Shoreface facies (USF) at the base. This is characterized by very well-sorted fine sands exhibiting predominantly plane lamination, which is consistent with its wave-dominated flow regime. The Central Shoreface facies (CSF) overlying the USF shows a high proportion of shells, a broad range of sediment sizes (fine sand to pebbles), and marked diversity in sedimentary structures. Graded storm beds, tidal cross-strata, mud-sand couplets, and horizontal to subhorizontal lamination are common. Swaley/hummocky (?) strata may also occur in its Distal subfacies. The sequence is topped by the Lower Shoreface facies (LSF). It consists mostly of bioturbated fine sands, although graded beds do occur. Distal sabfacies sands are consistently finer, better sorted, and more positively skewed than their Proximal counterpart. Anomalies in both coastwise and cross-shore textural trends are noted. In contrast to most other coastal settings, both fair-weather (wave- and tidal-current) and storm deposits are likely to be represented in subequal proportion in the rock record.     

Phylogenetic analysis of Toxicodendron （Anacardiaceae） and its biogeographic implications on the evolution of north temperate and tropical intercontinental disjunctions

Toxicodendron is a genus in the Rhus complex of Anacardiaceae with a disjunct distribution between eastern Asia and North America, extending to southeastern Asia and the neotropics. Nuclear （internal transcribed spacer, external transcribed spacer, and NIA-i3） and chloroplast （ndhF and trnL-F） sequences were used to construct phylogenetic relationships of Toxicodendron. Phylogenetic analysis of these data strongly support Toxieodendron as a monophyletic group distinct from other genera of the Rhus complex, and the phylogeny does not fully corroborate classification at the sectional level. Two temperate disjunct lineages were detected, one from section Toxicodendron and the other between the eastern North American Toxicodendron vernix and the eastern Asian Toxicodendron vernieifluum. Their divergence times were estimated to be 13.46 （7.95-19.42） and 7.53 （2.76-12.86） mya, respectively. The disjunction between section Griffithii （taxa from warm temperate to tropical Asia） and Toxieodendron striatum （from the neotropics） was supported and their divergence time was estimated to be 20.84 （11.1 6-30.52） mya in the early Miocene. Our biogeographic results and the paleontological data support the Bering land bridge as the most likely route to explain the temperate disjunctions, yet the tropical disjunction in Toxicodendron seems to be best explained by the North Atlantic land bridge hypothesis.     

Ontogenetic change of bone microstructures and its ethological implication in Champsosaurus (Diapsida,Choristodera)

Compact cortex in a Champsosaurus (Diapsida, Choristodera) femur is ontogenetically replaced with extensively developed cancellous bone. This histological shift, together with retention of calcified cartilage to late ontogenetic stage, was previously considered to show that adult champsosaurs were more adapted to aquatic environments than juveniles. However, the new histological examination reveals the nearly amedullar condition of a juvenile femur consisting of thick periosteal cortex and less cancellous bone tissue and the amedullar but more porous condition of adult femora. This likely demonstrates that the femoral inner structure of the juvenile is denser than those of the adults, and therefore, juveniles were more aquatic. It is suggested that morphological variations between two sympatric species of Champsosaurus reflect sexual dimorphism in a single species and limb bones with more robust morphology, showing better terrestrial adaptation for nesting on land, belong to females. The similarity of gross limb bone morphology between juveniles and inferred adult males indicates aquatic habitats for juveniles, coincident with the new interpretation of bone microstructures. No differences are, however, recognised in femoral microstructure between inferred sexes in adults. The possibly lowered density of femur in adults is considered as an adaptation to increase the mobility in water.     

Latitude,tree species diversity and the metabolic theory of ecology

The latitudinal diversity gradient (LDG) has been known for over a century, but its origin remains poorly understood. Because both latitude and species richness are broadly related to temperature, environmental temperature has been proposed as a driver of the LDG. Recently, Wang et al. (2009, Proceedings of the National Academy of Sciences USA, 106 ,13388–13392) used datasets compiled from tree distributions in eastern Asia and North America to compare the species richness?temperature relationship between the two regions at several spatial scales and framed their analyses in the context of the metabolic theory of ecology. Here, we show that their datasets lack comparability between eastern Asia and North America and that some aspects of their analyses probably biased their results, casting doubt on some of their conclusions.     

Pattern and process in the geographical ranges of freshwater fishes

North American freshwater fishes were studied to determine whether they displayed the same relationships between log (geographical range size) and log (body size) and the same pattern of range shape as found among North American birds and mammals. The forces that produce these patterns were also investigated. The log (geographical range size) : log (body size) relationship was analysed for 121 North American freshwater fish species. Thirty‐two imperilled species were compared with 89 non‐imperilled species to determine if the overall relationship could result from differential extinction. Range geometries were analysed, within and among habitat guilds, to determine if general patterns could be detected. The log (geographical range size) : log (body size) pattern among freshwater fish species was triangular and qualitatively similar to that found for North American birds and mammals. The results suggest that below a minimum geographical range, the likelihood of extinction increases dramatically for freshwater fishes and that this minimum range size increases with body size. The pattern of fish species’ range shapes differs from that found for other North American vertebrate taxa because, on average, fish possess much smaller ranges than terrestrial species and most fish species’ geographical ranges extend further on a north–south axis than on an east–west axis. The log (geographical range size) : log (body size) pattern reveals that fish species’ geographical ranges are more constrained than those of terrestrial species. The triangular relationship may be caused by differential extinction of species with large bodies and small geographical ranges as well as higher speciation rates of small‐bodied fish. The restricted geographical ranges of freshwater fishes gives them much in common with terrestrial species on oceanic islands. Range shape patterns within habitat guilds reflect guild‐specific historical and current ecological forces. The overall pattern of range shapes emerges from the combination of ecologically different subunits.