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The Bees of Portugal (Hymenoptera: Apoidea: Anthophila)
Abstract and Figures
The present paper provides a list of records, both from literature and from recent collections, of 680 bee species from mainland Portugal. Up to 1996 there were remarkably few published records of Portuguese bees, consisting of only around 325 species.
However, in the last 15 years or so there has been a considerable amount of collecting by numerous Hymenopterists, mainly in the Algarve, as well as inspection of unpublished material housed in museums and private collections. This effort has added over 300 species to the list. Further recording is needed, especially in the north. It is thought that the total number of bee species in Portugal could be as high as 700.
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... Low differentiation in COI analyses, combined with difficulties associated with the morphological separation of females of P. nanum and P. scapulare, and, in some cases, between females of P. nanum and P. stigmaticorne, resulted in the unresolved identification of some specimens and an ambiguous understanding of the geographic distributions of these species. In particular, all previous records of P. nanum on the Iberian Peninsula, including our own, were female specimens (Baldock et al., 2018). To further investigate the relationships among these taxa, and in an effort to identify female specimens important to the clarification of geographical distributions of these species, UCE data were obtained for 13 specimens representing P. nanum, P. scapulare, P. palestinicum and P. stigmaticorne. ...
... Distribution: Abkhazia, Albania, Armenia, Austria, Azerbaijan, Belgium, Bulgaria, Crimea, Croatia, Czech Republic, France (including Corsica), Georgia, Germany, Greece, Hungary, Italy (including Sardinia and Sicily), Kazakhstan, Malta, Moldova, the Netherlands, Romania, Russia (European part, Urals), Serbia, Slovakia, Slovenia, Switzerland, Turkey (European part), Turkmenistan and Ukraine (Fig. 6A). The single female specimen mentioned from Portugal in Baldock et al. (2018) was examined and determined to be a male P. stigmaticorne. Introduced in the USA. ...
The Palaearctic complex of anthidiine bees closely related to Pseudoanthidium scapulare has long been a source of unresolved taxonomic and systematic issues. Until now, the number of species in the complex and their geographical distributions were largely unclear, thus complicating the compilation of accurate species checklists and hindering conservation efforts. In order to address these issues, we use morphology and mitochondrial cytochrome c oxidase subunit I (COI) sequences, combined with a thorough examination of the relevant literature and type material, to delimit species within this complex, assign names to species and clarify geographical ranges. An unexpected result was that a certain number of morphologically distinct taxa exhibited low levels of genetic divergence at the COI locus, resulting in species paraphyly. A set of ultra-conserved elements (UCEs) was also sequenced in order to further investigate relationships among these taxa. One morphologically distinct species was also paraphyletic using UCE data, hinting at recent species divergences and genetic exchange at zones of contact between morphologically well-differentiated taxa. The results of our study reveal the presence of ten species in this complex, including a previously overlooked species for western continental Europe. A complete diagnosis of the males and females of these species is provided, as are maps detailing the geographic distributions of each. An illustrated identification key to the males and females of each species is presented. Two new species are described, Pseudoanthidium kaspareki sp. nov. and P. rozeni sp. nov. New synonymy is established for several species and Pseudoanthidium palestinicum and P. tropicum are raised to species level. The new combination, Icteranthidium floripetum comb. nov. is also established. Lectotypes are designated for the following species: Anthidium eversmanni, A. floripetum, A. frontale, A. karakalense, A. nanum and A. reptans. Previously unpublished lectotype designations are published here for A. sinuatum and A. tenellum.
... Label information from Ian Cross reports that a male specimen was collected at an aggregation of Melitturga caudata Pérez, 1879 (Andrenidae), on the sand near empty snail shells. Chrysis phryne has been reported to attack Osmia (Allosmia) melanura Morawitz, 1871 (see Pauli et al. (2019) (Baldock et al. 2018). We suggest that O. rufohirta is the likely host of C. crossi, though this must be confirmed through direct observations. ...
Background
DNA barcoding technologies have provided a powerful tool for the fields of ecology and systematics. Here, we present a part of the InBIO Barcoding Initiative Database: contribution to the knowledge on DNA barcodes of cuckoo wasps (Hymenoptera, Chrysididae) dataset representing 144 specimens and 103 species, covering approximately 44% of the Iberian and 21% of the European fauna. The InBIO Barcoding Initiative (IBI – DNA Barcoding Portuguese terrestrial invertebrate biodiversity) aims to fill the barcoding gap for the terrestrial invertebrate taxa. All DNA extractions are deposited in the IBI collection at CIBIO, Research Center in Biodiversity and Genetic Resources and specimens are deposited in the University of Mons collection (Belgium) and in the Natur- Museum in Lucerne (Switzerland).
New information
This dataset increases the knowledge on the DNA barcodes and distribution of 102 species of cuckoo wasps. A total of 52 species, from 11 different genera, were new additions to the Barcode of Life Data System (BOLD), with DNA barcodes for another 44 species added from under-represented taxa in BOLD. All specimens have their DNA barcodes publicly accessible through the BOLD online database. Nine cuckoo wasp species are newly recorded for Portugal. Additionally, two new species for science are described: Chrysis crossi Rosa, sp. nov. from southern Portugal and Hedychridium calcarium Rosa, sp. nov. from eastern Spain. Several taxonomic changes are proposed and Hedychrum rutilans Dahlbom, 1845 is found to consist of two different taxa that can be found in sympatry, Hedychrum rutilans s. str. and Hedychrum viridaureum Tournier, 1877 stat. nov. Stilbum westermanni Dahlbom, 1845 stat. nov. is confirmed as distinct from Stilbum calens (Fabricius, 1781), with the latter species not confirmed as present in Iberia; barcoded Stilbum material from Australia is distinct and represents Stilbum amethystium (Fabricius, 1775) sp. resurr.; Portuguese material identified as Hedychridium chloropygum Buysson, 1888 actually belongs to Hedychridium caputaureum Trautmann & Trautmann, 1919, the first confirmed record of this species from Iberia. Philoctetes parvulus (Dahlbom, 1845) is confirmed to be a synonym of Philoctetes punctulatus (Dahlbom, 1845). Chrysis lusitanica Bischoff, 1910 is confirmed as a valid species. Chrysis hebraeica Linsenmaier, 1959 stat. nov. is raised to species status.
... The tribe Anthidiini displays a wide variety of nesting behaviours, including nesting underground, using various types of cavities and building exposed nests (Michener 2007;Litman et al. 2016;Westrich 2018). Nesting in shells in this tribe has been recorded in only four Palaearctic species of Rhodanthidium (Erbar and Leins 2017;Baldock et al. 2018;Westrich 2018;Romero et al. 2020) and two Afrotropical species of Afranthidium Gess 2008, 2014). ...
Some species of two tribes (Anthidiini and Osmiini) of the bee family Megachilidae utilize empty gastropod shells as nesting cavities. While snail-nesting Osmiini have been more frequently studied and the nesting biology of several species is well-known, much less is known about the habits of snail-nesting Anthidiini. We collected nests of four species of the genus Rhodanthidium (R. septemdentatum, R. sticticum, R. siculum and R. infuscatum) in the Czech Republic, Slovakia, Catalonia (Spain) and Sicily (Italy). We dissected these nests in the laboratory and documented their structure, pollen sources and nest associates. The four species usually choose large snail shells. All four species close their nests with a plug made of resin, sand and fragments of snail shells. However, nests of the four species can be distinguished based on the presence (R. septemdentatum, R. sticticum) or absence (R. siculum, R. infuscatum) of mineral and plant debris in the vestibular space, and the presence (R. septemdentatum, R. infuscatum) or absence (R. sticticum, R. siculum) of a resin partition between the vestibular space and the brood cell. Rhodanthidium septemdentatum, R. sticticum and R. siculum usually build a single brood cell per nest, but all R. infuscatum nests studied contained two or more cells. For three of the species (R. siculum, R. septemdentatum and R. sticticum) we confirmed overwintering in the adult stage. Contrary to R. siculum, R. septemdentatum and R. sticticum do not hide their nest shells and usually use shells under the stones or hidden in crevices within stone walls. Nest associates were very infrequent. We only found two R. sticticum nests parasitized by the chrysidid wasp Chrysura refulgens and seven nests infested with pollen mites Chaetodactylus cf. anthidii. Our pollen analyses confirm that Rhodanthidium are polylectic but show a preference for Fabaceae by R. sticticum.
... It is estimated that Portugal is home to more than one thousand pollinating insect species, including a variety of bee, hoverfly, butterfly, and flower beetle species [16,23,24]. As of 2018, 680 distinct bee species have been collected and catalogued in Portugal [25]. Within the River Minho area alone, 200 distinct species were catalogued for a smaller scale study on pollination services [16]. ...
As urbanization and agriculture increase worldwide, habitats and food sources for wild
pollinators are often fragmented or destroyed. As wild pollinators contribute both resilience and variety to agricultural fields, it is desirable to implement land management practices that preserve their well-being and ability to contribute to food production systems. This study evaluates continental Portugal for its change in suitability to host bee’s pollinator species (Apis mellifera) from 1990 to 2018. It uses the InVEST crop pollination modeling tool and CORINE Land Cover, as well as parameterization to produce pollinator abundance and supply maps. These are generalized to municipality boundaries to provide actionable insights to farmers and policymakers and strengthen land management practices. It finds that the potential for pollination services is growing, with
averages of both pollinator abundance and supply indices improving by 8.76% across the continental territory in 28 years. The study results are validated using another pollination index derived from a study that is based on expert opinion and field sampling in a sub-region of Portugal. This method of aggregation of model results and comparison of the percent difference by administrative boundary has the potential to better inform both policymakers and farmers about the pollination potential on a local level, as well as inspire interventions for future productivity.
Revisionary taxonomic studies of bees from the Old World Mediterranean basin are hindered both by the apparent absence of type material for many taxa and a lack of genetic resources. The discovery of important type materials in combination with the generation of novel DNA barcodes (Cytochrome Oxidase I) has allowed cryptic diversity within the widespread taxa Nomiapis bispinosa (Brullé, 1832) and Systropha planidens Giraud, 1861 to be clarified. Nomiapis bispinosa actually consists of three distinct taxa: Nomiapis bispinosa s. str. from Morocco and Iberia to Central Asia, Nomiapis rufiventris (Spinola, 1838) spec. resurr. from Morocco to Egypt, including Sicily and Nomiapis paulyi spec. nov. from Portugal and Spain. A lectotype is designated for Nomia rufiventris Spinola, 1838. Lectotypes are designated for Nomia bispinosa Brullé, 1832 and Nomia albocincta Lucas, 1849, and type material for Nomia perforata Lucas, 1849 is clarified; both Nomiapis albocincta and Nomiapis perforata are synonymised syn. nov. with Nomiapis rufiventris. A lectotype is designated for Nomia ruficornis Spinola, 1838, and this taxon is confirmed as a synonym of Nomiapis bispinosa. Systropha planidens also consists of three distinct taxa: S. planidens from Central Europe to Iran and the European part of Russia, S. grandimargo Pérez, 1905 spec. resurr. from Portugal, Spain, and France, and S. anatolica Warncke, 1977 stat. nov. from Turkey, Syria, and northern Israel. A lectotype is designated for Systropha planidens Giraud, 1861. Systropha chrysura Pérez, 1905 is synonymised syn. nov. with S. grandimargo. These findings illustrate the extent to which our understanding of the taxonomy of Mediterranean bees remains incomplete.
The Hylaeus gibbus species-group is a notoriously difficult group, whose nomenclature is currently confusing. To clarify the taxonomy of this species group, the most problematic types of the gibbus species-group were examined. The identity of Hylaeus pictus (Smith, 1853) is unclear. The results suggest that two species are grouped under this name in Western Europe and in North Africa: H. pictus and H. purpurissatus (Vachal, 1895), stat. rev. Twelve synonymies are established: Hylaeus praenotatus Förster, 1871 and Prosopis bicarinata Pérez, 1903, n. syn. of Hylaeus gibbus Saunders, 1850; Prosopis nigripes Pérez, 1903, synonym of Hylaeus confusus Nylander, 1852; Prosopis morawitzi Pérez, 1903, Prosopis pereziana Cockerell, 1906, Prosopis pereziana Schulz, 1906 and Prosopis punctiscuta Alfken, 1909, n. syn. of Hylaeus distinguendus Morawitz, 1876; Prosopis stigmorhina Pérez, 1895, Prosopis stigmorhina var. basalis Pérez, 895, Hylaeus denisonae Cockerell, 1931, Hylaeus denisonae ifranensis Cockerell, 1931 and Hylaeus psilurus Cockerell, 1938, n. syn. of Hylaeus purpurissatus (Vachal, 1895). Lectotypes are designated for Prosopis picta Smith, 1853, Hylaeus praenotatus Förster, H. psilurus Cockerell and Prosopis purpurissata Vachal.
• In farmland landscapes worldwide, there are pervasive trends for either intensification or abandonment. Intensification is a widely recognised driver of wild bee declines, but little is known about the consequences of land abandonment, though it involves major habitat shifts from fallows, pastures and meadows, to shrublands and forests.
• Focusing on a Mediterranean landscape, we investigated long-term changes in wild bee communities during secondary vegetation succession after land abandonment. We used a space-for-time substitution approach, sampling plants and wild bees on five successional stages, from grasslands, through shrublands, to oak woodlands.
• We recorded 2721 bees, representing over 150 species. Grasslands had the highest wild bee abundance and high diversity of oligolectic species. Moving through successional stages, there was a rapid decline in bee abundance combined with a progressive accumulation of rare species, resulting in maximum diversity of the overall community, ground nesters, polyleges, and oligoleges in oak woodlands. Bee diversity was positively related to plant taxonomic and phylogenetic diversity. There was high turnover in community composition along the succession, with species occurring in grasslands being largely absent from later successional stages, and vice versa. From 21 indicator species, 17 were associated with grasslands, including the only threatened species.
• Our results suggest that a mosaic of habitats at different successional stages, particularly grasslands and oak woodlands, are necessary to maximise the diversity and abundance of wild bees at the landscape scale. Sustained management is thus needed under land abandonment to retain early-successional herbaceous habitats and to ensure woodland regeneration and protection.
Several studies recently reported that specialized (oligolectic) bees, which collect pollen from few host plants, use, besides visual cues, specific volatiles to find their hosts. Generalist (polylectic) bees, on the other hand, likely have to recognize a wider range of volatiles because they forage on many plant species. Bee antennal sensory equipment may thus be under selection to optimize plant host recognition. This selection may have led to variation in sensory equipment morphology with diet specialization (lecty). We tested if lecty correlates with antennal morphology and abundance of the main olfactory/gustatory sensilla (sensilla trichoidea (ST), placoidea (SP), sensilla basiconica (SB)) in the genus Andrena (Hyme-noptera: Andrenidae). Across 24 species, and after having controlled for body size, we found polylectic species to have a longer and narrower flagellomer F 9 (the one with highest abundance of sensilla), and to have a greater ST density on F 9 , compared with oligolectic species. Neither SP density nor SB number varied with lecty. A cluster analysis furthermore depicted groups of species that reasonably reflect diet specialization. Our results are in line with the previously observed lower number of glomeruli in the brain of oligolectic, compared with polylectic, bees. A formal correction for phylogeny is necessary to confirm our preliminary conclusion that pollen diet specialization has driven the morphology of the peripheral sensory system in this bee genus.
Andrena is a large genus of bees primarily distributed across the Holarctic. Despite their abundance in temperate regions, the pollen diets of many Nearctic Andrena remain incompletely resolved. The pollen diets of 50 species of Andrena found in Michigan were characterized using light microscopy. Twenty-four species (48%) were classified as pollen specialists, collecting pollen from one botanical family. The remaining 26 species (52%) were classified as pollen generalists, collecting from many botanical families. Andrena species fell into three broad foraging groups: (1) spring-flying species foraging predominantly from woody plants; (2) spring-flying species specializing on herbaceous ephemerals; and (3) summer-flying species specializing on herbaceous prairie plant species. Species of Nearctic Andrena specialized on pollens from Asteraceae, Geraniaceae, Hydrophyllaceae and Montiaceae or avoided them almost entirely. Botanical families that hosted specialized bees showed a higher variation in utilization by the Andrena community than botanical families without specialists. In contrast, Palaearctic species showed little temporal partitioning and low variation in the utilization of different botanical families. This pattern of pollen use supports previous findings that pollens from certain botanical families cannot be used as a food source without physiological adaptations, but that this phenomenon is more pronounced in the Nearctic Andrena fauna. ADDITIONAL KEYWORDS: Apoidea-Asteraceae-evolution-host plant specialization-oligolecty-pollen mixing-polylecty-solitary bee.
The osmiine bee species Hoplitis mucida is considered to consist of two subspecies with H. mucida mucida (Dours, 1873) ranging from northwestern Africa to Israel and Jordan and H. mucida stecki (Frey-Gessner, 1908) occurring in southwestern Europe and Sicily. The discovery of nests of H. mucida in Morocco and Tunisia revealed striking differences in the nesting biology of the two subspecies. In North Africa, females construct fully exposed, cake-like nests of mud on the flat surface of rocks and stones containing 8–12 vertically oriented brood cells, rendering these nests unique among osmiine bees regarding both nesting site and nest architecture. In contrast, in Europe females build their few-celled mud nests inside small rock cavities. This discrepancy in the nesting biology is paralleled by considerable morphological differences between the two subspecies suggestive of a long geographical isolation. Due to these biological and morphological differences, we propose to elevate the European subspecies H. mucida stecki to species rank.
The present study is an update to the first Red List of European Bees published in 2014. The additional records are based on (i) comprehensive review of literature; (ii) new data provided by bee specialists in response to the publication of the first Red List; (iii) new developments in taxonomy of European bees including description of new cryptic species; (iv) new specimens from recent field collections. While the first Red List included a list of 1965 wild bee species with 75 genera, we found 86 additional species, while two more genera have been erected (Seladonia and Vestitohalictus), giving an updated total of 2051 species and 77 genera. The authors discuss the artificial framework of the study considered by International Union for Conservation of Nature (IUCN) to produce the first Red List and they propose the more meaningful West Palaearctic biogeographical region. For this whole region, a first estimation gives 3408 wild bee species in 105 genera. The next taxa have been erected from subspecies to species status: Andrena (Euandrena) limosa Warncke, 1969, stat. n., Andrena (Plastandrena) oligotricha Mavromoustakis, 1952, stat. n.
Camptopoeum (Camptopoeum) baldocki spec. nov., a new European panurgine bee species is described and diagnosed. It is currently known only from saltmarshes along the southern coast of the central and eastern Algarve, Portugal. Observations and analysis of scopal pollen loads suggest narrow oligolecty on the similarly halophilous Frankenia laevis (Frankeniaceae). In addition, the allotype male of the Portuguese endemic Flavipanurgus fuzetus Patiny is described and the pollen preferences of Flavipanurgus are reviewed with the addition of new data from Portugal. As a genus, Flavipanurgus species appear to be narrowly oligolectic on a range of flowers from the botanical families Caryophyllaceae, Cistaceae and Crassulaceae.
RESUMEN El trabajo actual trata de los integrantes ibéricos de la familia Melittidae. Incluye la lista sinonímica de los taxones que componen esta fauna, así como, en cada una de las especies estudiadas, la distribución geográfica, el material revisado necesario para la discusión y, por lo tanto, para la confirmación o rechazo de las citas bibliográficas existentes hasta el presente, de igual modo que se pormenorizan los nuevos registros, tanto procedentes de capturas recientes como de material depositado en diferentes instituciones. La fauna ibérica de Melittidae está constituida por 3 subfamilias: Melittinae, Macropidinae y Dasypodinae; 3 géneros: Melitta, con 11 especies, Macropis, con 2 es-pecies, y Dasypoda, con 10 especies, que hacen un total de 23, correspondientes a 25 subespecies.
ABSTRACT Contribution to the knowledge of Iberian Melittidae. This paper deals with Iberian Melittidae. In every species, besides its geographical distribution, it includes the literature records that were mistaken or doubtful in order to confirm or correct them. The studied specimens, the new records and a checklist are included too. Three subfamilies belong to Iberian Melittidae: Melittinae, Macropidinae and Dasypodinae; 3 genera: Melitta, with 11 species, Macropis, with 2 species and Dasypoda, with 10 species, It results a total of 25 subspecies, belonging to 23 species.
Extreme specialization is a common phenomenon in antagonistic biotic interactions but it is quite rare in mutualistic ones. Indeed, bee specialization on a single flower species (monolecty) is a questioned fact. Here, we provide multiple lines of evidence on true monolecty in a solitary bee (Flavipanurgus venustus, Andrenidae), which is consistent across space (18 sites in SW Iberian Peninsula) and time (three years) despite the presence of closely related congeneric plant species whose flowers are morphologically similar. The host flower (Cistus crispus, Cistaceae) is in turn a supergeneralist, visited by at least 85 insect species. We uncover ultraviolet light reflectance as a distinctive visual cue of the host flower, which can be a key mechanism because bee specialization has an innate basis to recognize specific signals. Moreover, we hypothesized that a total dependence on an ephemeral resource (i.e. one flower species) must lead to spatiotemporal matching with it. Accordingly, we prove that the bee’s flight phenology is synchronized with the blooming period of the host flower, and that the densities of bee populations mirror the local densities of the host flower. This case supports the ‘predictable plethora’ hypothesis, that is, that host-specialization in bees is fostered by plant species providing predictably abundant floral resources. Our findings, along with available phylogenetic information on the genus Cistus, suggest the importance of historical processes and cognitive constraints as drivers of specialization in bee-plant interactions.
Changes in agricultural practice across Europe and North America have been associated with declines in wild bee populations. Bee diet breadth has been associated with sensitivity to agricultural intensification, but much of this analysis has been conducted at the categorical level of generalist or specialist, and it is not clear to what extent the level of generalisation within generalist species is also associated with species persistence. We used pollen load analysis to quantify the pollen diets of wild solitary bees on 19 farms across southern England, UK. A total of 72 species of solitary bees were recorded, but only 31 species were abundant enough to allow for formal diet characterisation. The results broadly conformed to existing literature with the majority of species polylectic and collecting pollen from a wide range of plants. Pollen load analysis consistently identified pollens from more plant species and families from each bee species than direct observation of their foraging behaviour. After rarefaction to standardise pollen load sample sizes, diet breadth was significantly associated with frequency of occurrence, with more generalist bees present on more farms than less generalist bees. Our results show that the majority of bee species present on farmland in reasonable numbers are widely variable in their pollen choices, but that those with the broadest diet were present on the greatest number of farms. Increasing the diversity of plants included in agri-environment schemes may be necessary to provide a wider range of pollen resources in order to support a diverse bee community on farmland.
Two new species of small carpenter bees are described from the West Palaearctic region: Ceratina zwakhalsi from South-East Turkey (compared with C. acuta Friese, C. callosa (Fabricius) and C. chalybea Chevrier) andCeratina verhoeffi from Morocco (compared with C. saundersi Daly and C. maghrebensis Daly). Distribution maps are given for all the threated species. Ceratina saundersi is recorded for the first time from Portugal, the Lampedusa island (south of Sicily), the Italian peninsula and Lybia. The occurence of C. saundersi in Sardinia and Spain is confirmed.
The loss of key forage plants and a narrow pollen diet have both been implicated in the decline of wild bees over the past 70 years. These ideas have been studied extensively in recent years in bumblebees (Bombus spp.), but have rarely been investigated in other bee groups, due in part to a lack of detailed ecological data for many species of wild bee. Chambers (1968) extensively documented pollen preferences in bees from the genus Andrena collected in Bedfordshire, UK, during the 1940s, before the period of rapid agricultural intensification following the Second World War. This extensive dataset allows for the importance of diet breadth in modifying the response of bees to environmental change to be investigated in a common and widespread but understudied genus of bees. We compared the pollen preferences of a suite of Andrena species collected in the 1941-1949 period with a similar suite of Andrena species collected between 1985 and 2016. Relative diet breadth was consistent across studied species between the historical and contemporary period. However, dietary composition changed with a shift away from Rosaceae and towards Brassicaceae. The reduction in the collection of Rosaceae pollen was more pronounced in spring-flying than summer-flying Andrena, which may be due to a reduction in the length and quality of hedgerow habitats and hence spring-flowering woody Rosaceae present in the countryside, and an increase in the availability of the mass-flowering crop oilseed rape (Brassica napus). Both historical and contemporary diet breadth were significant positive predictors of how frequently Andrena species are encountered on contemporary farmland, with those species with a relatively wider diet present on the majority of farms. These findings support the idea that inherent differences in diet breadth mediate the ability of bee species to respond to changes in resource availability resulting from agricultural intensification. A more detailed understanding of species-level characteristics can help improve our understanding of why seemingly similar species respond very differently to environmental change.