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An Annotated Checklist of Invasive Species of the Phylum Arthropods and Chordates in Panama

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28 June 2024

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28 June 2024

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Abstract
Invasive species are one of the five main causes of biodiversity loss, along with habitat destruction, overexploitation, pollution, and climate change. Represent one of the first barriers to overcome in ecological conservation programs, since are difficult to control and eradicate. Due to the lack of records of invasive exotic species in Panama, this research is necessary to identify and register the documented groups of invasive species of the Chordates and Arthropod groups in Panama. This exhaustive search of invasive species was carried out in different bibliographic databases, electronic portals and scientific journals which addressed the topic at a global level. The results indicate that approximately 133 invasive exotic species of the Arthropoda and Chordata Phylum have been reported in Panama. Of the 133 species, 49 species belong to the Arthropoda Phylum and 84 species belong to the Chordate Phylum. The Panamanian economic activity could promote the introduction of alien species in this country. The study provides the first list of invasive exotic chordate and arthropod species reported for the Republic of Panama.
Keywords: 
Invasive species
;  
introduced species
;  
biodiversity
;  
Central America
;  
tropical forest
;  
pest
;  
Panama
Subject: 
Biology and Life Sciences  -   Biology and Biotechnology

1. Introduction

Invasive species are one of the five main causes of biodiversity loss, along with habitat destruction, overexploitation, pollution, and climate change. An invasive species is understood to be an exotic species that establishes itself in a natural, semi-natural ecosystem or habitat, is an agent that induces changes and affects native biodiversity [1]. The process of invasion is a progressive phenomenon that doesn´t have to be unidirectional [2]. It means, not all introduced species (exotic) will become naturalized, nor all naturalized ones will become invasive. Neither exist a constant proportion of species that go from one phase to another [2].
The problems associated with invasive species are summarized in three main aspects [3]: (1) the big economic costs and ecological impacts caused by many invasive species; (2) the increase in the number of introduced species that become invasive and in consequence the problems associated with these species; and (3) the needs to include the invasive species in any ecological study, taking in mind that the current nature dynamics are governed by many invasive species [3].
Globally, 90% of vertebrate and plant introductions are intentional and the remaining 10% are accidental [4]. Invasive species are one of the first barriers to overcome in ecological conservation programs, since are difficult to control and eradicate, mainly due to their reproductive strategies combined with their dispersal, establishment, and persistence [3].
Invasive alien species can be found in all groups of organisms [5]. After plants, arthropods and chordates include the largest number of invasive exotic species reported for Latin America and the Caribbean [4].
The animal groups which pertain to the Phylum Arthropoda contain the largest number of species known today [6]. Some species transmit diseases or are vectors of them, and some can affect crops and many serve as natural indicators and pollinators. In addition, a few of these species play the role of primary, secondary consumers or decomposers [7].
Most morphologically and physiologically complex organisms are found in the Chordate Phylum, they have a high diversity of ecological niches, with notable terrestrial and aquatic environment adaptations [7]. In terms of ecosystemic level, the introduction of mammals can affect the food chain function, generating a cascade effect in the composition and abundance of invasive predator, herbivore, and plant species, as well as nutrient cycles [8]
Panama is the twenty-eighth country in the world with the greatest biological diversity [9]. However, in proportion to its size it ranks tenth. Panama has a higher number of known vertebrate animals than any other country in Central America or the Caribbean and more bird species than the United States and Canada together [9]. In addition, it has 3.5% of the plants with flowers and 7.3% of the fern and related species in the world [10]. Due to its geographical position, high land, and maritime connectivity, as well as the existence of the Panama Canal, Panama has helped biological invasions [10]. According to [11], the new locks used in the expansion of the Panama Canal allow passage of ships that are 71 meters longer and 16 meters wider, allowing more mobilization of exotic species within its ballast waters or attached to the ship hull [10].
According to the IV Panama National Biodiversity Report, an estimate of approximately 324 exotic species were introduced in Panama, the majority are plants. In addition, the introduction of exotic species as pets has increased. Among the exotic species used as pets are birds, reptiles, and mammals [10]; however, it is not known how many of these exotic species have become invasive species. Due to the lack of records of invasive exotic species in Panama, this research is necessary to help answer how many invasive species of the group of chordates and arthropods have been registered in Panama.
This research will provide the first list of invasive species in the Panamanian territory of two important groups of organisms: arthropods and chordates. This research will serve as preliminary data on the sources of income and will provide information for future research and plans to prevent the impact of those species.

2. Materials and Methods

2.1. Study Area

Panama is geographically located at 7° 12' 07'' and 9° 38'46'' north latitude and 77° 09' 24'' and 83° 03' 07'' west Neotropical Region longitude. Panama is in the central part of the American continent, in the most eastern and south part of Central America. It limits to the north with the Caribbean Sea, to the south with the Pacific Ocean, to the east with Colombia and to the west with Costa Rica [9].
With a land area of 75,416.6 km2, administratively composed of ten provinces, three indigenous regions (Kuna Yala, Emberá, Ngäbe Buglé) with province status, two regions (Kuna de Madungandí and Kuna de Wargandí) with the level of municipal government, and 623 townships throughout the country [12].
Panama bases its economy on four activities, one of them being the logistics industry, which is fundamentally based on the movement of cargo from all over the world [13]. This movement that develops through ports, airports, railways, and the Panama Canal is one of the activities recognized as a means of introducing invasive species [14], in addition to the already known Zona del Canal that during that time allowed the intentional introduction of several species [10].

2.2. Study Selection

The study was carried out through an exhaustive information review, in the following sources; Global Invasive Species Database: http://www.iucngisd.org/gisd/species.php?sc=965, Invasive Species Specialist Groupup IUCN/SSC, Invasive Species Specialist Group (ISSG), Invasive Species Compendium (CABI) https:// www.cabi.org/isc/datasheet/108530#tolistOfSpecies. In bibliographic databases such as Web of Science (WOS), Science Direct, Scielo, PubMed, Redalyc, Dimensions and Google Scholar. Using the Publish or Perish software, the URLs resulting from the searches were downloaded in csv format, which was worked in Excel to filter those records that contained the mixture of words (invasive species, Panama alien species, invasive species in Panama, invasive exotic species in Panama, Panama invasive species). Additionally, thorough search was carried out in specialized journals on the subject, such as: BioInvasions Records (Reabic), Biological Invasions (Springer), Revista Bioinvasiones, Check List the Journal of biodiversity data (PENSOFT), as well as the journals published in Panama that are found on the ABC Platforms (SENACYT) such as National Resources and SIBIUP of the University of Panama.
The search was done under the keywords of: invasive species, exotic species, invasive species in Panama, invasive exotic species in Central America, invasive species of insects in Panama, invasive species in America, New records + Panamá, Pest + Panama, introduced species + Panamá, “Alien species”, and filtered by the word Panama, New records + Panama, Pest + Panama, Introduced species + Panama.
Due to the limited literature published on the subject for Panama, the country's specialists from the different groups under study were asked for a list of the species considered invasive in the country. We visited Environment Ministry bibliography and reviewed the subject related, the Aquatic Resources Authority of Panama, the Panama Canal Authority, the Herbarium of the Panama University and the Directorate of Plant Health of the Ministry of Agricultural Development.
Other sources of information used were: Fifth National Report of Panama to the Convention on Biological Diversity [15]; Invasives in Mesoamerica and the Caribbean (International Union for nature and natural resources conservation 2002); National Biodiversity Strategy and Action Plan 2018-2050 [12], Ministerio de Desarrollo Agropecuario database of invasive species 2019.
The information obtained was organized in tables for easier understanding. A database of the species was created (in cases where the information was obtained) with the following variables: report date (year), province, altitude, place of origin. This allowed us to answer the research question and meet one of the objectives. Also to make easy the access to information on invasive species of the Phylum Arthropoda and Chordate, another two tables were created, one that included the taxonomic identification of the groups and the reference according to the article or database that was obtained for the species, and the last one that contained pets limited distribution under official control in Panama Republic.

3. Results

Through the collection of information, the results indicate that approximately 134 invasive exotic species of the Arthropoda and Chordata Phylum have been reported in Panama. Of the 133 species, 49 species belong to the Arthropoda Phylum and 84 species belong to the Chordate Phylum, indicated in taxonomy table 1.
Of the 49 species belonging to the Arthropoda Phylum, 36 species belonged to the Insecta class, three to the Arachnida class, nine to the Malacostraca class and one to the Maxillopoda class. Of the 84 species of the Chordate Phylum, 42 belonged to the Actinopterygii class, 20 to the Ascidiacea class, eight species to the Reptilia class, six to the bird class, and four to the Amphibia and Mammalia class.
In the Arthropoda Phylum, 18 species belong to the Coleoptera order, being the most abundant of the groups in this Phylum followed by Decapoda with nine species, Hymenoptera with six species, Diptera, Hemiptera with five each group. Acarida with two species, and one species in the Lepidoptera, Blattodea, Mesostigmata and Cephalobaenida orders.
In the Chordate Phylum, 11 belong to the Stolidobranchia order, 10 to Cichliformes, eight to the Cypriniformes order, seven species belong to Squamata order, six species belong to the Perciformes, Cyprinodontiformes, and Aplousobranchia orders, four to the Passeriformes and Anura orders, three to the Characiformes and Phlebobranchia orders, two in the Rodentia, Carnivora, Salmoniformes and Gobiiformes orders, and one species for the Columbiformes, Pelecaniformes, Carangiformes, Bleniformes, Acanthuriformes, Scorpaeniformes, Siluriformes and Testudines orders.
The three most abundant families of the Arthropoda Phylum are: Curculionidae (14 spp.), Formicidae (4 spp.), Tephritidae (3 spp.). For the Chordata Phylum the three most abundant families are: Cichlidae (10 spp.), Styelidae (8 spp.) and Cyprinidae (7 spp.).
There´s one case we called “Eye Attention” for Rachycentron canadum (Linnaeus, 1766), were the data located it in the Pacific zone of the Panama Republic, but have never seem again in other Panama aquatic ecosystem.
Table 1. Exotic invasive species taxonomy list from the Panama Republic.
Table 1. Exotic invasive species taxonomy list from the Panama Republic.
Species Family Order Class Phylum Reference
Hypothenemus hampei Ferrari, 1867 Curculionidea Coleoptera Insecta Arthropoda [26]
Coccotrypes advena Blandford 1894 Curculionidae Coleoptera Insecta Arthropoda [27]
Ambrosiodmus obliquus LeConte, 1878 Curculionidae Coleoptera Insecta Arthropoda [27]
Coptoborus ricini Eggers, 1932 Curculionidae Coleoptera Insecta Arthropoda [27]
Xyleborus bispinatus Eichhoff, 1864 Curculionidae Coleoptera Insecta Arthropoda [27]
Xylosandrus morigerus Blandford, 1894 Curculionidae Coleoptera Insecta Arthropoda [27]
Xylosandrus compactus Eichhinoff, 1876 Curculionidae Coleoptera Insecta Arthropoda [27]
Cryptocarenus seriatus Eggers, 1933 Curculionidae Coleoptera Insecta Arthropoda [27]
Coccotrypes vulgaris Eggers, 1923 Curculionidae Coleoptera Insecta Arthropoda [27]
Dendrocranulus tardus Blandford, 1896 Curculionidae Coleoptera Insecta Arthropoda [27]
Scolytopsis puncticollis Blandford, 1896 Curculionidae Coleoptera Insecta Arthropoda [27]
Xylosandrus crassiusculus Motschulsky, 1866 Curculionidae Coleoptera Insecta Arthropoda [28]
Xyleborinus exiguus Walker, 1859 Curculionidae Coleoptera Insecta Arthropoda [27]
Rhynchophorus palmarum Linneus, 1758 Curculionidae Coleoptera Insecta Arthropoda [29]
Euoniticellus intermedius Reiche, 1849 Scarabaeidae Coleoptera Insecta Arthropoda [30]
Harmonia axyridis Pallas, 1773 Coccinellidae Coleoptera Insecta Arthropoda [31]
Callosobruchus phaseoli Gyllenhal, 1833 Chrysomelidae Coleoptera Insecta Arthropoda [32]
Saperda candida Fabricus, 1787 Cerambycidae Coleoptera Insecta Arthropoda [33]
Brachyplatys subaeneus Westwood, 1837 Plastaspidae Hemiptera Insecta Arthropoda [34]
Diaphorina citri Kuwayama, 1908 Psyllidae Hemiptera Insecta Arthropoda [35]
Pseudacysta perseae Heideman, 1908 Tingidae Hemiptera Insecta Arthropoda [36]
Aulacaspis yasumatsui Takagi, 1977 Diaspididae Hemiptera Insecta Arthropoda [37]
Aphis spiraecola Patch, 1914 Aphididae Hemiptera Insecta Arthropoda [38]
Apis mellifera Linneus, 1758 Apidae Hymenoptera Insecta Arthropoda [39]
Tapinoma melanocephalum Fabricius, 1793 Formicidae Hymenoptera Insecta Arthropoda [33]
Paratrechina longicornis Motshulsky, 1863 Formicidae Hymenoptera Insecta Arthropoda [33]
Monomorium floricola Jerdon, 1851 Formicidae Hymenoptera Insecta Arthropoda [33]
Quadrastichus erythrinae Kim, 2004 Eulophidae Hymenoptera Insecta Arthropoda [40]
Nylanderia fulva Mayr, 1862 Formicidae Hymenoptera Insecta Arthropoda [41]
Ceratitis capitata Weidemann, 1824 Tephritidae Diptera Insecta Arthropoda [42]
Anastrepha obliqua Macquart, 1835 Tephritidae Diptera Insecta Arthropoda [43]
Anastrepha grandis Macquart, 1846 Tephritidae Diptera Insecta Arthropoda [44]
Aedes aegypti Linnaeus, 1762 Culicidae Diptera Insecta Arthropoda [45]
Aedes albopictus Skuse, 1895 Culicidae Diptera Insecta Arthropoda [45]
Tuta absoluta Meyrick, 1917 Gelechiidae Lepidoptera Insecta Arthropoda [46]
Blattella germanica Linnaeus, 1767 Ectobiidae Blattodea Insecta Arthropoda [47]
Brevipalpus phoenicis Geijskes, 1936 Tenuipalpidae Acarida Arachnida Arthropoda [48]
Brevipalpus californicus Banks, 1904 Tenuipalpidae Acarida Arachnida Arthropoda [49]
Dermanyssus gallinae De Geer, 1778 Dermanyssidae Mesostigmata Arachnida Arthropoda [50]
Rhithropanopeus harrisii Gould, 1841 Panopeidae Decapoda Malacostraca Arthropoda [51]
Elamenopsis kempi Chopra y Das, 1930 Hymenosomatidae Decapoda Malacostraca Arthropoda [52]
Procambarus clarkia Girard, 1852 Cambaridae Decapoda Malacostraca Arthropoda [44]
Penaeus monodon Fabriciud, 1798 Penaeidae Decapoda Malacostraca Arthropoda [44]
Cherax tenuimanus Smith, 1912 Parastacidae Decapoda Malacostraca Arthropoda [44]
Macrobrachium amazonicum Heller, 1862 Palemonidae Decapoda Malacostraca Arthropoda [53]
Macrobrachium rosenbergii De Man, 1879 Palemonidae Decapoda Malacostraca Arthropoda [62]
Goniopsis cruentata Latrelle, 1803 Grapsidae Decapoda Malacostraca Arthropoda [53]
Pachygrapsyus gracilis de Saussure, 1857 Grapsidae Decapoda Malacostraca Arthropoda [53]
Raillietiella frenatus Riley & Self, 1981 Cephalobaenidae Cephalobaenida Maxillopoda Arthropoda [54]
Mus musculus Linnaeus, 1758 Muridae Rodentia Mammalia Chordata [55]
Rattus norvegicus Berkenhout, 1769 Muridae Rodentia Mammalia Chordata [55]
Herpestes auropunctatus Hodgson, 1836 Herpestidae Carnivora Mammalia Chordata [33]
Herpestes javanicus Hilaire, 1818 Herpestidae Carnivora Mammalia Chordata [33]
Columba livia Gmelin, 1789 Columbidae Columbiformes Aves Chordata [33]
Bubulcus ibis Linnaeus, 1758 Ardeidae Pelecaniformes Aves Chordata [33]
Mimus gilvus Viellot, 1808 Mimidae Passeriformes Aves Chordata [33]
Passer domesticus Linnaeus, 1758 Passeridae Passeriformes Aves Chordata [56]
Quiscalus mexicanus Gmelin, 1788 Icteridae Passeriformes Aves Chordata [55]
Sicalis flaveola Linnaeus, 1766 Thraupidae Passeriformes Aves Chordata [56]
Rachycentron canadum Linnaeus, 1766 Rachycentridae Carangiformes Actinopterygii Chordata [57]
Oreochromis niloticus Linnaeus, 1758 Cichlidae Cichliformes Actinopterygii Chordata [58] , [59]
Oreochromis aureus Steindachner, 1864 Cichlidae Cichliformes Actinopterygii Chordata [58]
Oreochromis mossambicus Peters, 1852 Cichlidae Cichliformes Actinopterygii Chordata [58]
Oreochromis urolepis Norman, 1912 Cichlidae Cichliformes Actinopterygii Chordata [58]
Coptodon rendalli Boulenger, 1897 Cichlidae Cichliformes Actinopterygii Chordata [58]
Cichla ocellaris Bloch & Schneider, 1801 Cichlidae Cichliformes Actinopterygii Chordata [33]
Cichla monoculus Agassiz, 1831 Cichlidae Cichliformes Actinopterygii Chordata [58] , [59]
Parachromis managuensis Günther, 1867 Cichlidae Cichliformes Actinopterygii Chordata [60] , [59]
Astronotus ocellatus Agassiz, 1881 Cichlidae Cichliformes Actinopterygii Chordata [59]
Mesonauta festivus Heckel, 1840 Cichlidae Cichliformes Actinopterygii Chordata [59]
Eleotris melanosoma Bleeker, 18531859 Eleotridae Gobiiformes Actinopterygii Chordata [61]
Butis koilomatodon Bleeker, 1849 Butidae Gobiiformes Actinopterygii Chordata [61]
Lepomis macrochirus Rafinesque, 1819 Centrarchidae Perciformes Actinopterygii Chordata [62]
Lepomis microlophus Günther, 1859 Centrarchidae Perciformes Actinopterygii Chordata [63]
Micropterus salmoides Lacepede, 1802 Centrarchidae Perciformes Actinopterygii Chordata [63] , [59]
Pomoxis annularis Rafinesque, 1818 Centrarchidae Perciformes Actinopterygii Chordata [61]
Pomoxis nigromaculatus Lesueur, 1829 Centrarchidae Perciformes Actinopterygii Chordata [61] , [59]
Lepomis humilis Girard, 1858 Centrarchidae Perciformes Actinopterygii Chordata [59]
Omobranchus punctatus Valenncienes, 1836 Blenniidae Bleniiformes Actinopterygii Chordata [61]
Sciaenops ocellatus Linnaeus, 1766 Sciaenidae Acanthuriformes Actinopterygii Chordata [44]
Oncorhynchus mykiss Walbaum, 1792 Salmonidae Salmoniformes Actinopterygii Chordata [33] , [59]
Salmo trutta Linnaeus, 1758 Salmonidae Salmoniformes Actinopterygii Chordata [63]
Pterois volitans Linnaeus, 1758 Scorpaenidae Scorpaeniformes Actinopterygii Chordata [64]
Cirrhinus chinensis Günther, 1868 sin C. molitorella Cyprinidae Cypriniformes Actinopterygii Chordata [44]
Cirrhinus molitorella Valenciennes, 1844 Cyprinidae Cypriniformes Actinopterygii Chordata [44]
Ctenopharyngodon Idella Valenciennes, 1844 Cyprinidae Cypriniformes Actinopterygii Chordata [61]
Cyprinus carpio Linnaeus, 1758 Cyprinidae Cypriniformes Actinopterygii Chordata [33] , [59]
Hypophthalmichthys molitrix Valenciennes, 1844 Cyprinidae Cypriniformes Actinopterygii Chordata [61]
Hypophthalmichthys nobilis Richardson, 1845 Cyprinidae Cypriniformes Actinopterygii Chordata [33]
Mylopharyngodon piceus Richardson, 1846 Cyprinidae Cypriniformes Actinopterygii Chordata [44]
Ictiobus cyprinellus Valenciennes, 1844 Catostomidae Cypriniformes Actinopterygii Chordata [44]
Gambusia holbrooki Girard, 1859 Poeciliidae Cyprinodontiformes Actinopterygii Chordata [59]
Xiphophorus helleri Heckel, 1848 Poeciliidae Cyprinodontiformes Actinopterygii Chordata [59] , [65]
Gambusia affinis Baird & Girard, 1853 Poeciliidae Cyprinodontiformes Actinopterygii Chordata [65]
Gambusia nicaraguensis Günther, 1866 Poeciliidae Cyprinodontiformes Actinopterygii Chordata [65]
Poecilia reticulata Peters, 1859 Poeciliidae Cyprinodontiformes Actinopterygii Chordata [65]
Xiphophorus variatus Meek, 1904 Poeciliidae Cyprinodontiformes Actinopterygii Chordata [65]
Ictalurus punctatus Rafinesque, 1818 Ictaluridae Siluriformes Actinopterygii Chordata [44]
Colossoma macropomum Cuvier, 1816 Serrasalmidae Characiformes Actinopterygii Chordata [61] , [59]
Piaractus brachypomus Cuvier, 1818 Serrasalmidae Characiformes Actinopterygii Chordata [44]
Piaractus mesopotamicus Holmberg, 1887 Serrasalmidae Characiformes Actinopterygii Chordata [44]
Hemidactylus brookii Gray, 1845 Gekkonidae Squamata Reptilia Chordata [66]
Hemidactylus frenatus Dúmeril & Bibron, 1836 Gekkonidae Squamata Reptilia Chordata [66] , [33]
Hemidactylus mabouia Moreau de Jonnes, 1818 Gekkonidae Squamata Reptilia Chordata [66] , [33]
Hemidactylus turcicus Linnaeus, 1758 Gekkonidae Squamata Reptilia Chordata [66] , [67]
Lepidodactylus lugubris Dúmeril & Bibron, 1836 Gekkonidae Squamata Reptilia Chordata [66] , [68]
Anolis sagrei Cocteauin Dúmeril & Bibron, 1837 Polychrotidae Squamata Reptilia Chordata [69]
Sphaerodactylus argus Gosse, 1850 Sphaerodactylidae Squamata Reptilia Chordata [66] , [62]
Trachemys scripta elegans Wied, 1838 Emydidae Testudines Reptilia Chordata [33]
Eleutherodactylus antillensis Reinhardt & Lütken, 1863 Eleutherodactylidae Anura Amphibia Chordata [70] , [71]
Eleutherodactylus johnstonei Barbour, 1914 Eleutherodactylidae Anura Amphibia Chordata [70]
Eleutherodactylus planirostris Cope, 1862 Eleutherodactylidae Anura Amphibia Chordata [72]
Rana catesbeiana Shaw, 1802 Ranidae Anura Amphibia Chordata [44]
Ascidia incrassata Heller, 1878 Ascidiidae Phlebobranchia Ascidiacea Chordata [73]
Ascidia sydneiensis Stimpson, 1855 Ascidiidae Phlebobranchia Ascidiacea Chordata [73]
Phallusia nigra Savigny, 1816 Ascidiidae Phlebobranchia Ascidiacea Chordata [73]
Botrylloides nigrum Herdman, 1886 Styelidae Stolidobranchia Ascidiacea Chordata [73]
Botryllus planus Van Name, 1902 Styelidae Stolidobranchia Ascidiacea Chordata [73]
Polyandrocarpa anguinea Sluiter, 1898 Styelidae Stolidobranchia Ascidiacea Chordata [73]
Polyandrocarpa sagamiensis Tokioka, 1953 Styelidae Stolidobranchia Ascidiacea Chordata [73]
Polyandrocarpa zorritensis Van Name, 1931 Styelidae Stolidobranchia Ascidiacea Chordata [73]
Styela canopus Savigny, 1816 Styelidae Stolidobranchia Ascidiacea Chordata [73]
Symplegma brakenhielmi Michaelsen, 1904 Styelidae Stolidobranchia Ascidiacea Chordata [73]
Symplegma rubra Monniot, 1972 Styelidae Stolidobranchia Ascidiacea Chordata [73]
Herdmania pallida Heller, 1878 Pyuridae Stolidobranchia Ascidiacea Chordata [73]
Microcosmus exasperatus Heller, 1878 Pyuridae Stolidobranchia Ascidiacea Chordata [73]
Pyura vittata Stimpson, 1852 Pyuridae Stolidobranchia Ascidiacea Chordata [73]
Didemnum perlucidum Monniot F, 1983 Didemnidae Aplousobranchia Ascidiacea Chordata [73]
Didemnum psammatodes Sluiter, 1895 Didemnidae Aplousobranchia Ascidiacea Chordata [73]
Diplosoma listerianum Milne Edwards, 1841 Didemnidae Aplousobranchia Ascidiacea Chordata [73]
Lissoclinum fragile Van Name, 1902 Didemnidae Aplousobranchia Ascidiacea Chordata [73]
Trididemnum orbiculatum Van Name, 1902 Didemnidae Aplousobranchia Ascidiacea Chordata [73]
Polyclinum constellatum Savigny, 1816 Polyclinidae Aplousobranchia Ascidiacea Chordata [73]
Based on the information obtained, we can say that the province with the highest record of invasive species introduction in the Panama Republic is the Panama province (79 records), followed by the Colon (16 records), Chiriqui province (ten records), Veraguas (six records), Cocle province (four records), West Panama and Bocas del Toro (three records), Herrera (two records), Los Santos, San Blas and Darien province (one record), (Table 2).
The four continents with more record of invasive species origin that had been reported in Panama are: Asia (22 records), South America (21 records), Africa (15 records) and North America (14 records) (Table 2).
Among the species reported as invasive for the Panama Republic, nine species are under official control, all belonging to the insect class, causing damage to cucurbit crops, tomato, citrus, coffee, and species of the Arecaceae family (Table 3).

4. Discussion

According to our results from the Arthropoda and Chordate Phylum, there are an estimated 133 invasive species for the Panama Republic. Comparing with the information from other Iberoamerican countries we found that for the countries that are part of the Andean Community (Colombia, Ecuador, Venezuela, Peru and Bolivia) 227 invasive exotic species have been identified, mostly plants (92 species), insect pests (61 species) and vertebrates (30 species) [16]. In Costa Rica 235 invasive species have been reported from all groups of organisms [17]. For the Dominican Republic, 192 invasive species have been reported, were 38 fish, 4 amphibians, 8 reptiles, 13 birds and 13 mammals [18]. While in other latitudes such as the Iberian Peninsula (Spain, Portugal and Andora) 100 invasive species have been recorded [19].
When comparing the number of invasive species based on the surface area of each country (Andean Community and Dominican Republic) in which this study is addressed, we found that for the Panama Republic the number is higher: Panama (134 species), compared to the Andean Community (166 species), Dominican Republic (138 species).
Panama has 75,416.6 km2 area regard to the countries of the Andean community; Colombia (1,142 million km2), Ecuador (256,370 km2), Venezuela (916,445 km2), Peru (1,285 million km2), Bolivia (1,099 million km2) and the Dominican Republic with an area of 48,442 km2, which indicates that the Panama Republic, even though it is a small country compared to those previously mentioned, it has quite a considerable number of invasive species in the territory.
Considering economic activity of the Dominican Republic based in beaches tourism [20], Colombia in the free market (exchange of goods and services between people and companies through monetary transactions) [21], Ecuador and Venezuela in oil revenues [22,23], Peru in the exploitation, processing and export of natural resources, mainly mining, agricultural and fishing [24], Bolivia in the natural gas [25], and since the Panama Republic is a country that bases its economy as a logistics and transportation center, commercial and financial center, we can assume that this is the reason why the constant influx of invasive species is reflected so high compared to the others countries.

5. Conclusions

The results indicate that approximately 133 invasive exotic species of the Arthropoda and Chordata Phylum have been reported in Panama. Of the 133 species, 49 species belong to the Arthropoda Phylum and 84 species belong to the Chordate Phylum. When comparing the number of invasive species based on the surface area of each country, in which this study is addressed, we found that for the Panama Republic the number is higher: This research can serve as a basis for future studies in the country that are of vital importance for pest control and recommendations for the complete collection of information. This research benefits the wealth of the country and therefore the care of our resources, and provides more information to entities seeking management plans to solve problems caused by the invasion of species.
Within the limitations of the work, it must also be considered that it is possible that some species are not included due to the dispersion of the information or that they have not been considered invasive for some researchers because they may correspond to natural dispersal patterns, like is the case of the Coyote.

Author Contributions

Conceptualization, E.M and D.R.; methodology, E.M and D.R; research, E.M, H.G, R.F, L.R, Y.A, O.L and D.R; Writing-preparation of original draft, E.M and D.R; Writing-revisions and editing, E.M, H.G, R.F, L.R, Y.A, O.L and D.R; acquisition of funds, E.M. All authors have read and approved this version of the manuscript for publication.

Funding

This research was funded by the University of Panama, the Panamanian National Research System (SNI) and through SENACYT DDCCT economic subsidy contract No. 004-2023.

Institutional Review Board Statement

Not applicable.

Informed Consent Statement

Not applicable.

Data Availability Statement

Not applicable.

Acknowledgments

To Darío Luque who provided us with the database of invasive exotic species of MIAMBIENTE; to Mgtr. Mirna Samaniego from STRI who look for information in the Web Of Science; To Pedro Méndez, Ricardo Moreno, Jorge García and Chelina Bastista for assisting with the names of the invasive species for their respective groups; to Mrs. Digna in the MIAMBIENTE documentation section for providing the books on invasive species; and to the Marine Resources Authority of Panama for offering information on fish introduced in Panama.

Conflicts of Interest

The authors declare no conflicts of interest.

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Table 2. Information about: invasive species, province, district, site, year, species altitude was found for the first time in Panama Republic, place of origin, affectation.
Table 2. Information about: invasive species, province, district, site, year, species altitude was found for the first time in Panama Republic, place of origin, affectation.
Species Province District Site IntroductionBREAKYear SiteBREAK altitude Belonging BREAKContinet
Hypothenemus hampei Chiriqui Chiriqui Border with Costa Rica 2007 Africa
Coccotrypes advena Panama, Colon Panama, Colón Panama Canal 2007 Asia
Ambrosiodmus obliquus Panama
Coptoborus ricini Panama
Xyleborus bispinatus Panama
Xylosandrus morigerus Panama
Xylosandrus compactus Panama
Cryptocarenus seriatus Panama
Coccotrypes vulgaris Panama
Dendrocranulus tardus Panama
Scolytopsis puncticollis Panama
Xylosandrus crassiusculus Panama Gatun Barro Colorado 2004 10 m Asia
Xyleborinus exiguus Panama Gatun Barro Colorado 2019 10 m Asia
Rhynchophorus palmarum Panama Panama, Colon Panama
Euoniticellus intermedius Chiriqui Renacimiento Chiriqui 2015 Africa
Harmonia axyridis Colon Colon Centro Regional Colon 2014 Europa
Callosobruchus phaseoli Panama Arraijan Las villas de Arraijan 2022 Neotropic
Saperda candida
Brachyplatys subaeneus Herrera Chitre Costa Pacífica 2016 Asia
Diaphorina citri Bocas del Toro Bocas del Toro Bocas del Toro 2016 Asia
Pseudacysta perseae Los Santos Santa Maria Peninsula de Azuero 2017 28 a 49 Europe
Aulacaspis yasumatsui Panama Panama Balboa 2011-2021 4m-50m Asia
Aphis spiraecola Cocle Penonome, Ola Cocle 2002 180 a 440 South America
Apis mellifera Panama Panama Panama Republic Africa
Tapinoma melanocephalum
Paratrechina longicornis
Monomorium floricola
Nylanderia fulva Chiriqui Chiriqui Chiriqui
Ceratitis capitata Chiriqui Chiriqui Chiriqui 1963 South America
Anastrepha obliqua Cocle Cocle Cocle 2011 14 a 1550
Anastrepha grandis Darien La Palma Darien 2009 20 South America
Aedes aegypti Panama Panama Panama 1985 Africa
Aedes albopictus Panama Panama Panama 2003 Asia
Tuta absoluta Panama Panama Panama
Quadrastichus erythrinae Panama Panama Panama Canal 2018 South America
Blattella germanica Panama Panama Panama Africa
Brevipalpus phoenicis
Brevipalpus californicus
Dermanyssus gallinae Herrera Ocu Las Guabas 2018
Rhithropanopeus harrisii Panama, Colon Gatun Lake 1969 Noth America
Elamenopsis kempi Panama, Colon Gatun Lake 2007 Africa
Procambarus clarkii Panama Panama 1987
Penaeus monodon Panama Panama Panama North America
Cherax tenuimanus Panama Panama Panama Oceania
Macrobrachium amazonicum Panama, Colon Gatun Lake
Macrobrachium rosenbergii Panama 1981 Asia
Goniopsis cruentata Panama, Colon Gatun Lake
Pachygrapsyus gracilis Panama, Colon Gatun Lake
Raillietiella frenatus Panama
Mus musculus Panama Panama Panama 1973 Europe, Asia
Rattus norvegicus Panama Panama Panama Asia
Herpestes auropunctatus Panama Panama Panama Asia
Herpestes javanicus
Columba livia Panama Panama Panama 1608 Europe, Africa, Asia
Quiscalus mexicanus Panama Panama Panama North America
Passer domesticus Panama Panama Panama 2019 Europe
Sicalis flaveola Panama Panama Panama South America
Mimus gilvus Panama
Bubulcus ibis Panama
Rachycentron canadum Panama Gulf of Panama 2015 South America
Oreochromis niloticus Chiriqui, Veraguas, Panama, Colon Fortuna, La Yeguada, Gatun, Alajuela, Bayano and rivers 1976 Africa
Oreochromis aureus Panama Panama Panama Canal 1976 Northwest Africa
Oreochromis mossambicus Panama Gatún, Alajuela Panama Canal 1940 Southeast Africa
Oreochromis urolepis Panama Panama Panama Canal 1979 Southeast Africa
Coptodon rendalli Panama Panama Panama Canal 1977 Africa, Middle east
Cichla ocellaris Panama, Colon Gatun Lake 1967 South America
Cichla monoculus Panama, Colon, Veraguas Gatun, Alajuela y La Yeguada 1969 South America
Parachromis managuensis Chirqui, Veraguas, Panama, Colon Fortuna, La Yeguada, Gatun, Alajuela and rivers 1973 Central America (Atlantic Slope)
Astronotus ocellatus Panama, Colon Gatun Lake, Alajuela 1991 South America
Mesonauta festivus Panama, Colon Gatun Lake 1998-1999 South America
Eleotris melanosoma Panama Panama Panama Africa, Asia, Indo-Pacific
Lepomis macrochirus Chiriqui, West Panama, Panama Lagunas de Volcan y San Carlos, Las Cumbres 1955 North America
Lepomis microlophus Panama Panama Panama North America
Micropterus salmoides Chiriqui Lagunas de Volcan 1955 North America
Pomoxis annularis Panama Panama Panama North America
Pomoxis nigromaculatus West Panama La Laguna Laguna de San Carlos 1925-1935 North America
Omobranchus punctatus Panama Panama Panama Canal 2011 East Asia
Lepomis humilis West Panama La Laguna Laguna de San Carlos 1925-1935 North America
Sciaenops ocellatus Cocle Aguadulce El Salado 1987 North America
Oncorhynchus mykiss Chiriqui Chiriqui Viejo, Caldera, Boquete 1925 North America
Salmo trutta Panama Panama Panama Europe
Pterois volitans Colon Colon Panamanian caribbean 2008 Pacific ocean, Asia
Butis koilomatodon Panama Panama Miraflores locks 1973 Occidental Indo-Pacific
Cirrhinus chinensis Günther, 1868 sin C. molitorella Panama Panama Panama 1977 Asia
Cirrhinus molitorella Panama Panama Panama 1980 Asia
Ctenopharyngodon idella Panama Panama Panama 1978 Asia
Cyprinus carpio Chiriqui, Veraguas Fortuna, La Yeguada 1976 Europe, Asia
Hypophthalmichthys molitrix Panama Panama Panama 1978 Asia
Hypophthalmichthys nobilis Panama Panama Panama 1978 Asia
Mylopharyngodon piceus Panama Panama Panama East Asia
Ictiobus cyprinellus Veraguas Santiago Santiago 1987 Cuba
Ictalurus punctatus Panama Panama Panama 1981 North America
Gambusia holbrooki Panama, Colon Gatun Lake North America
Xiphophorus helleri Cocle, Panama Las Cumbres, Anton Valley 1950´s y 1970´s Central America (Atlantic Slope)
Gambusia affinis Panama Panama Panama
Gambusia nicaraguensis Panama Panama Panama
Poecilia reticulata Veraguas, Panama Santiago, Pacora, Chame, Las Cumbres, Cerro Azul 1910-1912 South America
Xiphophorus variatus Panama Panama Panama
Colossoma macropomum Panama Panama? Panama? Es La Yeguada, río Sereno y Alajuela 1980 South America
Piaractus brachypomus Panama Panama Panama 1980 South America
Piaractus mesopotamicus Panama Panama Panama 1986-1987 South America
Hemidactylus brookii Panama Panama Tocumen
Hemidactylus frenatus Panama Panama Chepo, Tocumen 2019 36 Tropical region, Indo-Pacific
Anolis sagrei Panama Panama Albrook 2018 Cuba
Hemidactylus mabouia Panama Panama Canal zone 1994 sub-Saharan Africa
Hemidactylus turcicus Panama Panama Canal Zone 1914
Lepidodactylus lugubris Not specific 2004 Siri Lanka-Africa
Sphaerodactylus argus Bocas del Toro, San Blas Bocas del Toro Bocas del Toro, San Blas 2002 Jamaica, Caribean island
Trachemys scripta elegans Panama Panama Panama United States
Eleutherodactylus antillensis Panama Panama Panama city 1950-1960 Puerto Rico, Virgin Island
Eleutherodactylus johnstonei Panama Panama Panama 1980
Eleutherodactylus planirostris Panama Panama Panama Cuba, Bahamas
Rana catesbeianus Panama Panama Panama North America
Ascidia incrassata Panama Panama Panama canal 2009 Indo Pacific
Ascidia sydneiensis Panama Panama Panama Australia-Oceania
Phallusia nigra Panama Panama Panama Indo Pacific
Botrylloides nigrum Panama Panama Panama Canal 2002 South America
Botryllus planus Panama Panama Panama Indo Pacific
Polyandrocarpa anguinea Panama Panama Panama Africa
Polyandrocarpa sagamiensis Panama Panama Panama Suth America
Polyandrocarpa zorritensis Panama Panama Panama Suth America
Styela canopus Colon Colon Panama canal Indo Pacific
Symplegma brakenhielmi Panama Panama Panama
Symplegma rubra Panama Panama Taboguilla Indo- Pacific
Herdmania pallida Bocas del Toro Colon Island Bocas del Toro South America
Microcosmus exasperatus Panama Panama Panama Colombia-South America
Pyura vittata Panama Panama Panama
Didemnum perlucidum Panama Panama Panama South America
Didemnum psammatodes Panama Panama Panama
Diplosoma listerianum Panama Panama Panama
Lissoclinum fragile Panama Panama Panama
Trididemnum orbiculatum Panama Panama Panama South America
Polyclinum constellatum Panama Panama Panama
Table 3. Information about: invasive species, province, district, site, year, species altitude was found for the first time in Panama Republic, place of origin, affectation.
Table 3. Information about: invasive species, province, district, site, year, species altitude was found for the first time in Panama Republic, place of origin, affectation.
Pests Class Species Host
South American cucurbit fly insecta Anastrepha grandis Macquart, 1846 Cucurbitaceae
Tomato moth insecta Tuta absoluta Meyrick, 1917 Tomato
Huanglongbing (HLB) insecta Diaphorina citri Kuwayama, 1908 Citrus
Mediterranean fly Insecta Ceratitis capitata Wiedemann, 1824 fruit plants (citrus, cucurbitaceae)
Oriental Trips palmi Insecta Thrips palmi Karny, 1990 Cucurbitaceae
Coffe berry borer Insecta Hypothenemus hampei Ferrari, 1867 Coffe
Citrus leprosis virus Insecta Brevipalpus californicus Banks, 1904 Citrus
Insecta Brevipalpus phoenicis Geijskes, 1936 Citrus
Red Ring Disease Insecta Rhynchophorus palmarum Linnaeus, 1758 Arecaceae
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