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Paleobotanical notes on mangrove-like plants of Puerto Rico/Notas paleobotanicas acerca de plantas parecidas a mangles en Puerto Rico/Notas paleobotanicas sobre plantas parecidas a mangues em Porto Rico.


Several new trace fossils (ichnofossils) of mangrove-like plants are reported for Puerto Rico; all these fossils are new records for Puerto Rico. The earliest root casts in Puerto Rico were dated Late Cretaceous (probably Santonian, 85.8-83.5Ma). Earlier paleobotanical studies in Tertiary-dated sites in Puerto Rico are also discussed. Oligocene and Miocene lignitic rocks, traces of amber, and trace fossils (possibly mangrove rhizoliths) also have been found in several geological formations of northern and southern Puerto Rico (e.g., Juana Diaz Formation and Ponce Limestone). Rhizoliths of Scaveola cf. plumieri (L.) Vahl has been collected from coastal Pleistocene eolianite terraces that occur in northern Puerto Rico. The Puerto Rican material is herein described and illustrated, and a brief paleobotanical discussion provided. To date, the data on ichnofossils of mangrove-like plants are too poor to speculate on such subjects as dispersal and island biogeography, especially in Puerto Rico. Most of the taxonomic information presented herein was collected from fragmentary surveys. However, the data herein suggest that the composition of fossilized mangrove-like plants is more complex than previously suspected. Therefore, it would seem important to continue studying such paleoenvironments, in order to contribute to the conservation and knowledge of the paleoecology of mangrove-like plants of Puerto Rico.

KEYWORDS / Mangrove / Paleobotany / Puerto Rico /


Se informa para Puerto Rico varios nuevos fosiles traza (icnofosiles) de plantas parecidas a los mangles; todos estos fosiles son nuevos registros para Puerto Rico. Los moldes de raices mas antiguos en Puerto Rico fueron datados del Cretacico Tardio (probablemente Santoniano, 85,8-83,5Ma). Tambien se discuten los estudios paleobotanicos previos en los yacimientos datados del Terciario en Puerto Rico. Las rocas ligniticas, trazas de ambar y fosiles trazas (probablemente rizolitos de mangle) del Oligoceno y Mioceno tambien se han encontrado en varias formaciones geologicas del norte y sur de Puerto Rico (e.g., la Formacion Juana Diaz y la Caliza Ponce). Los rizolitos de Scaveola cf. plumieri (L.) Vahl han sido colectados de las terrazas costeras de eolianita del Pleistoceno que ocurren al norte de Puerto Rico. El material puertorriqueno se describe y se ilustra y se proporciono una breve discusion paleobotdnica. Al presente, los datos sobre icnofosiles de plantas parecidas a los mangles es muy escasa como para especular sobre temas como la dispersion y biogeografia de la isla, especialmente en Puerto Rico. La mayor parte de la informacion taxonomica aqui presentada fue reunida de los reconocimientos fragmentados. Sin embargo, los datos sugieren que la composicion fosilifera de las plantas parecidas a los mangles es mas complejo que lo previamente sospechado. Por consiguiente, parece importante el continuar estudiando dichos paleoambientes para contribuir a la conservacion y el conocimiento de la paleoecologia de las plantas parecidas a los mangles de Puerto Rico.


Informa-se para Porto Rico, varios novos fosseis traca (icnofosseis) de plantas parecidas aos mangues; todos estes fosseis sao novos registros para Porto Rico. Os moldes de raizes mais antigos em Porto Rico foram datados do Cretaceo Tardio (provavelmente Santoniano, 85,8-83,5Ma). Tambem se discutem os estudos paleobotanicos previos nas jazidas datadas do Terciario em Porto Rico. As rochas ligniticas, tracas de ambar e fosseis tracas (provavelmente rizolitos de mangue) do Oligoceno e Mioceno tambem se tem encontrado em varias formacoes geologicas do norte e sul de Porto Rico (e.g., a Formacao Juana Diaz e a Caliza Ponce). Os rizolitos de Scaveola cf. plumieri (L.) Vahl tem sido recolhidos dos terracos costeiros de eolianitos do Pleistoceno que ocorrem ao norte de Porto Rico. O material porto-riquenho descrito e ilustrado proporcionou uma breve discussao paleobotanica. Atualmente, os dados sobre icnofosseis de plantas parecidas aos mangues sao muito escassos como para especular sobre temas como a dispersao e biogeografia da ilha, especialmente em Porto Rico. A maior parte da informacao taxonomica aqui apresentada foi reunida dos reconhecimentos fragmentados. No entanto, os dados sugerem que a composicao fossilifera das plantas parecidas aos mangues e mais complexa que o previamente suspeitado. Por conseguinte, parece importante continuar estudando ditos paleoambientes para contribuir a conservacao e o conhecimento da paleoecologia das plantas parecidas aos mangues de Porto Rico.


A mangrove forest is considered a dynamic ecotone (or transition zone) between terrestrial and marine habitats. In its simplest sense "mangrove", as used herein, encompasses a group of woody, halophylic plants that occurs along sheltered tropical and subtropical coastlines. Mangroves are derived from a variety of plant families and vary in their dependence upon littoral habitats. Mangrove forests are also referred to as mangrove swamps, tidal forests, tidal swamp forests or mangals. Caribbean mangroves range from 30[degrees]N (northern Florida) to 8[degrees]N (northwest Colombia) and from 59[degrees]W (north of Guiana) to 89[degrees]W (eastern Guatemala) (Tomlinson, 1986). Until recently, four mangrove species were known to occur in Puerto Rico (Lugo and Snedaker, 1974; Francis and Lowe, 2000; Little et al., 2001), the most widely distributed of which are the red mangrove (Rhizophora mangle L.), black mangrove (Avicennia germinans (L.) Stearns), white mangrove (Laguncularia racemosa (L.) Gaertn.), and buttonwood (Conocarpus erectus L.). Following Tomlinson (1986), only three species should be considered as "true mangroves" (A. germinans, R. mangle, and L. racemosa) in Puerto Rico. Conocarpus erectus is frequently considered a "true mangrove", but Tomlinson (1986) suggested that it is better regarded as a mangrove associate because it lacks of the biological features (pneumatophores and vivipary) which characterize true mangroves; furthermore, it occurs in inland communities. A compilation of 125 plants routinely found in mangrove forests, the introduction of a new variety of the mangrove-associated plant, C. erectus var. sericeus Griseb. (Combretaceae), unusual aerial roots in A. germinans (Avicenniaceae) were reported for Puerto Rico (Nieves-Rivera, 2005). The latter study produced an actualized checklist of scientific and common botanical names (English/ Spanish), including 3 true mangroves (A. germinans, L. racemosa and R. mangle), and 122 mangrove-associates for Puerto Rico.

The most common trace fossils, also known as ichnofossils or "Lebensspuren" ('living traces' in German) left by plant activity are root traces or casts (rhizoliths), which show the branching and irregularities of living root morphology. These casts also reveal plant behavior, by showing growth. Their preservation taphonomy was probably the result of early cementation around the original roots, followed by cementation of carbonate sand that filled the mold (Martin, 1996). The fossilized mangrove-like plants of Puerto Rico have been less studied. A total of 11 trace fossils (ichnofossils) of mangrove-associated plants are herein reported for Puerto Rico (Table I).

The earliest root casts in Puerto Rico were dated Late Cretaceous (probably Santonian, 85.8-83.5Ma; Santos, 1990, 1999). These plant ichnofossils (Figure 1a-e), along with scattered large, near-vertical trace fossils (Skolithos s. str.; Prothero and Schwab, 1996) are present on the top of bioturbated magnetite-rich units (magnetite lenses) of the oldest facies of the Cotui Limestone, Cabo Rojo-San German, in southwestern Puerto Rico (Santos, 1990, 1999). The location of the Cotui Limestone root casts is given on Table I and was at 25.3m elevation. Cretaceous root casts have been tentatively interpreted to have been caused by mangrove-like plants, e.g., Deltoidospora sp. (= Acrostichum) or the extinct Brevitricolpites sp., although recent palynological studies reported the origin of these two candidates in the Eocene (Tomlinson, 1986; Rull, 1998). Another possible candidate is the mangrove-associated palm Nypa, which nowadays is widely distributed throughout southwestern Asia and dates back to Late Cretaceous (Tomlinson, 1986; Rull, 1998). These casts (2-20mm wide), which Santos suggested that were derived from some mangrove-like plants, possess many anatomical traits also found in modern mangrove-associated plants (e.g., Acrostichum spp.). The original wood material was decomposed and no longer exists; however, the spaces were filled with soft sediment, easily removed by physical and biogeochemical mechanisms (Figure 1c-e). These features are characteristic of intertidal to shallow subtidal marine environments with high energy (active waves and currents) (Santos, 1990, 1999).


In the 1920s, Hollick (1928) began a paleobotanical survey in Puerto Rico, which was summarized in his Paleobotany of Porto Rico, based on his field collections of seeds, leaves, and wood macrofossils of various species, including mangrove species such as Rhizophora, in the gray shale walls of the Collazo and Guatemala Rivers of the San Sebastian Formation (SSF), of Oligocene in age (33.7-23.8Ma). These plant species were also collected in other localities around the island (Hollick, 1928). This author reported leaves of Rhizophora sp. (Rhizophora (sp.) doctrinalis Hollick; Figure 5b, plate 82 of Hollick, 1928) from station B (at the base of the first falls below the bridge) in the gray shales of the Collazo River, SSF, northwestern Puerto Rico. A type (YPM 27218), paratype (YPM 27196), and figure (YPM 27199) of R. (sp.) doctrinalis were deposited in the Yate Peabody Museum (Table I).

Palynological studies in Puerto Rico were conducted by Graham and Jarzen (1969) and Graham (1996, 2003), collecting at many of Hollick's original Tertiary surveyed sites, and adding new ones. Palynomorphs (pollen) of these plant species were collected in the shales and organic-rich silty limestone layers of the San Sebastian and Lares Formations, northwestern Puerto Rico. In their study, Graham and Jarzen (1969) obtained 165 palynomorphs; 44 were identified and 15 are unknown. Graham (1996, 2003) summarized Hollick's works and demonstrated that there is further need for paleobotanical studies in the region on the diversity and importance of the fossil plant record of Puerto Rico. Graham (1995) carried out palynological studies in the Caribbean, with emphasis on the diversification of the Gulf/Caribbean mangrove communities, especially before and after the appearance of the Isthmus of Panama in the Pliocene (5.3-1.8Ma; s. str. Graham, 1992).

Although fossil pollen of Avicennia has not been collected in Puerto Rico, this mangrove was the first to be found in the Late Miocene (11.2-5.3Ma) of the Caribbean (Graham, 1995), although Duke (1995) reported Avicennia in Early Miocene (23.8-16.4Ma). Avicennia pollen is also common in the Quaternary (1.8-0.01Ma) of Costa Rica, Panama, and in northern South America (Muller et al., 1987; Graham, 1995). Microfossils of Rhizophora first appeared in the Late Eocene (37.0-33.7Ma), Avicennia in Late Miocene, Laguncularia in the Pliocene (5.3-3.6Ma), and Conocarpus in the Quaternary (Graham, 1995).

More recent palynological surveys by Graham and Jarzen (1969) and Graham (1995, 1996, 2003), reported Puerto Rican Tertiary (Middle Oligocene) mangrove plant microfossils. Mangrove pollen included Rhizophora sp. and Pelliciera sp. (Graham and Jarzen, 1969; Graham, 1995, 1996, 2003). Palynomorphs of both mangrove species were collected in the light gray and gray shales of SSF. Both mangrove species are typical of coastal habitats with brackish or marine waters; however, Pelliciera rhizophorae Triana & Planchon is now limited to the Pacific coasts of Costa Rica, Panama, and both Pacific and Atlantic coasts of Colombia (Tomlinson, 1986).

Oligocene and Miocene lignitic rocks, traces of amber, and trace fossils (possibly mangrove rhizoliths) also have been found in several geological formations of northern and southern Puerto Rico (e.g., Juana Diaz Formation and Ponce Limestone s. str. Frost et al., 1983; MacPhee and Wyss, 1990; MacPhee and Iturralde-Vinent, 1995; Iturralde-Vinent, 2001; Nieves-Rivera, unpubl, data, 2002; Table I). The biostratigraphy of the (Holiday Inn) outcrop, which is part of the Ponce Limestone, is Late Miocene in age and is herein reported (Maria Ruiz-Yantin, pers. comm., 2004): "During Late Miocene times, this area was active tectonically, whereas the northern part of Puerto Rico was passive. The outcrop has four units. Unit 1 shows a lagoon environment and it is 1.2m in thickness. It is a wackestone composed mainly of the foraminifer Miosorites cf. americanus Seiglie & Grove, solitary corals, Pecten with original shells, and internal gastropod molds. Unit 2 represents a reef front environment and is 4.5m thick; it is a massive unit (packstone) containing solitary corals in growth position, colonial corals such as Diploria and Porites, unidentified burrows, Pecten, crabs remains, M. cf. americanus foraminifers, rhizoliths of mangrove origin (LACMIP Ponce, PR 17772), and internal gastropods molds; it shows a coral framework. Unit 3 represents a reef crest environment and is 4.5m thick; The corals are out of place and are not as well as cemented in the underlying unit; there are solitary corals, Montastrea annularis (Ellis & Solander), Porites porites (Pallas), brain corals that are probably Diploria; also, there are few gastropods, bivalves, and crabs remains; there is an erosion surface between unit 3 and 4. Unit 4 is 8.05m thick and shows a lagoon environment similar to Unit 1".

Pleistocene eolianite terraces (s.str. Taggart, 1992) that occur in southwestern Mona Island and northern Puerto Rico jut out from the coastline. The shoreline of Mona Island terraces sometime shows "dead mangrove roots" that protrude from crannies and fissures, and coral fossils (e.g., M. annularis, M. cavernosa L., Diploria sp., Acropora palmata Lamarck) are found everywhere (Hernandez-Avila, 1970; Taggart, 1992). In Punta Jacinto, located at Playa Jobos in Isabela, northern Puerto Rico, there is a typical example of aeolian fossilized dunes having plant root casts (rhizoliths). The Isabela rhizoliths were deposited in the Natural History Museum of Los Angeles County, Department of Invertebrate Paleontology, Los Angeles, California (LACMIP). Pleistocene plant rhizoliths (see LACMIP Isabela PR 17768) are possibly from a mangrove-like plant such as Scaveola cf. plumieri (L.) Vahl (Storrs L. Olson, pers. comm., 2005; Figure 2a-f; Table I). Similar rhizoliths have been found as reefs at Key Biscayne Bay in Florida (USA; Hoffmeister and Multer, 1965), in aeolian dunes of Hawaii (Olson and James, 1982), and San Salvador Island in the Bahamas (Martin, 1996).


To date, the data on ichnofossils of mangrove-like plants are too poor to speculate on such subjects as dispersal and island biogeography, especially in Puerto Rico. Most of the taxonomic information presented herein was collected from fragmentary surveys from Puerto Rico. However, the data herein suggest that the composition of fossilized mangrove-like plants is more complex than previously suspected. Therefore, it would seem important to continue studying such paleoenvironments, in order to contribute to the conservation and knowledge of the paleoecology of mangrove-like plants of Puerto Rico.


The author thanks Hernan Santos, Jorge Velez-Juarbe and Maria Ruiz-Yantin (Dept. of Geology, University of Puerto Rico at Mayaguez -UPRM-) for their help in locating the trace fossils; "Storrs L. Olson (Smithsonian Institution, Washington, D.C.) for his help on rhizoliths; Alan Graham (Missouri Botanical Garden, St. Louis, Missouri) for his aid on Table I; Juan A. Rivero (Dept. of Biology, UPRM) and Ariel E. Lugo (USDA, Institute of Tropical Forestry at Rio Piedras) for corrections to the text; and Peter Rocafort (Dept. of Marine Sciences, UPRM) for digitalization of figures. This project was supported by the University of Puerto Rico Alliance for the Graduate Education and the Professorate fellowship (Grant No NSF/AGEP-HRD # 0302696).

Received: 03/27/2006. Accepted : 02/01/2007.


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Angel M. Nieves-Rivera. B.S. in Biology, Inter American University of Puerto Rico, San German, Puerto Rico. M.S. in Biology, University of Puerto Rico, Mayaguez (UPRM). Puerto Rico. Ph.D. in Marine Sciences, UPRM, Puerto Rico. Address: Agriculture Specialist, U.S. Customs and Border Protection, Office of Field Operations, John F. Kennedy International Airport, Terminal 4, Jamaica, NY 11430, NY, USA. e-mail:

Taxa                                Accession number (1)

Mangrove-like root casts (IF)       --

Ilex sp. (PL)                       MO C-47,15; ESF H-19

Myrica sp. (PL)                     MO C-42,4; ESF N-43

Pelliciera sp. (PL)                 MO C-48, 1; ESF D-38,2

Rhizophora (sp.) doctrinalis

Hollick (IF)                        YPM 27218 (type)
                                    YPM 27196 (paratype)
                                    YPM 27199 (figure)
Rhizophora sp. (PL)                 MO C-47,1; ESF L-21,3

Tournefortia sp. (PL)               MO A-12,10; ESF U-34,4

Mangrove-like root casts (IF) *     --

Mangrove-like root casts (IF) *     LACMIP 17772

Scaveola cf. plumieri rhizoliths    LACMIP 17768
(IF) *

Mangrove-like root casts (IF)       --

Taxa                                    Coordinates

Mangrove-like root casts (IF)       8[degrees] 04.720'N,
                                    67[degrees] 05.312'W

Ilex sp. (PL)                       18[degrees] 19.998'N,
                                    66[degrees] 56.918'W

Myrica sp. (PL)                     18[degrees] 19.998'N,
                                    66[degrees] 56.918'W

Pelliciera sp. (PL)                 18[degrees] 19.998'N,
                                    66[degrees] 56.918'W

Rhizophora (sp.) doctrinalis        18[degrees] 19.998'N,
Hollick (IF)                        66[degrees] 56.918'W

Rhizophora sp. (PL)                 18[degrees] 19.998'N,
                                    66[degrees] 56.918'W

Tournefortia sp. (PL)               18[degrees] 17.413'N,
                                    66[degrees] 53.411'W

Mangrove-like root casts (IF) *     17[degrees] 58.412'N,
                                    66[degrees] 53.730'W

Mangrove-like root casts (IF) *     17[degrees] 58.947'N,
                                    66[degrees] 40.028'W

Scaveola cf. plumieri rhizoliths    18[degrees] 30.873'N,
(IF) *                              67[degrees] 04.501'W

Mangrove-like root casts (IF)       18[degrees] 04.854' N,
                                    67[degrees] 56.379'W

Taxa                                 Geological      Geologic
                                    Formation (2)    Time (3)

Mangrove-like root casts (IF)            CM             LC

Ilex sp. (PL)                            SC             MO

Myrica sp. (PL)                          SC             MO

Pelliciera sp. (PL)                      SC             MO

Rhizophora (sp.) doctrinalis             SC             MO

Hollick (IF)

Rhizophora sp. (PL)                      SC             MO

Tournefortia sp. (PL)                    SL             MO

Mangrove-like root casts (IF) *          BL             MO

Mangrove-like root casts (IF) *          PC             LM

Scaveola cf. plumieri rhizoliths         ED             PL
(IF) *

Mangrove-like root casts (IF)            EF             QT

Taxa                                References

Mangrove-like root casts (IF)       Santos (1990, 1999);

                                    Nieves-Rivera (2005)
Ilex sp. (PL)                       Graham and Jarzen (1969)

Myrica sp. (PL)                     Graham and Jarzen (1969)

Pelliciera sp. (PL)                 Graham and Jarzen (1969)

Rhizophora (sp.) doctrinalis        Hollick (1928);

Hollick (IF)                        Graham (1996)

Rhizophora sp. (PL)                 Graham and Jarzen (1969)

Tournefortia sp. (PL)               Graham and Jarzen (1969)

Mangrove-like root casts (IF) *     Nieves-Rivera (2005)

Mangrove-like root casts (IF) *     Nieves-Rivera (2005)

Scaveola cf. plumieri rhizoliths    Storrs L. Olson
(IF) *                              (pers. comm., 2005);
                                    Nieves-Rivera (2005)

Mangrove-like root casts (IF)       Hernandez-Avila (1970);
                                    Nieves-Rivera (2005)

* New record for Puerto Rico; ESF: England Slide Finder coordinates.

(1) Museum abbreviations. LACMIP: Natural History Museum of Los
Angeles County, Department of Invertebrate Paleontology, Los Angeles,
California; MO: Herbarium, Missouri Botanical Garden, Saint Louis,
Missouri; YPM: Yale Peabody Museum Herbarium, Yale University,

(2) Geological formations. BL: Barrio Luna, Juana Diaz Formation,
Guanica; CM: Cotui Mountains, Cotui Formation, Cabo Rojo-San German;
ED: Eolianite dunes, Punta Jacinto, Isabela; EF: Eolianite Formation,
Carabinero, Mona Island; PC: Ponce Cement Quarry, Ponce Limestone,
Ponce; SC: Salto Collazo, San Sebastian Formation, San Sebastian; SL:
Slope, junction of Roads PR-111 and PR-124, San Sebastian Formation,

(3) Geologic Time. LC: Late Cretaceous (Santonian, 85.8-83.5Ma); LM:
Late Miocene (11.2-5.3Ma); MO: Middle Oligocene (28.5Ma); PL:
Pleistocene (1.8-0.01Ma); QT: Quaternary (1.8Ma-10000 yr).
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